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2017-04-01 Phylogenetics, Biogeography, and Patterns of Diversification of GeckosAcross the Sunda Shelf with an Emphasis on the GenusCnemaspis (Strauch, 1887) Perry Lee Wood Brigham Young University

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BYU ScholarsArchive Citation Wood, Perry Lee, "Phylogenetics, Biogeography, and Patterns of Diversification of GeckosAcross the Sunda Shelf with an Emphasis on the GenusCnemaspis (Strauch, 1887)" (2017). All Theses and Dissertations. 7259. https://scholarsarchive.byu.edu/etd/7259

This Dissertation is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in All Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Phylogenetics,
Biogeography,
and
Patterns
of
Diversification
of
Geckos

Across
the
Sunda
Shelf
with
an
Emphasis
on
the
Genus

Cnemaspis
(Strauch,
1887)

Perry
Lee
Wood
Jr.

A
dissertation
submitted
to
the
faculty
of Brigham
Young
University in
partial
fulfillment
of
the
requirements
for
the
degree
of

Doctor
of
Philosophy

Jack
W.
Sites
Jr.,
Chair Byron
J.
Adams Seth
M.
Bybee L.
Lee
Grismer Duke
S.
Rogers

Department
of
Biology

Brigham
Young
University

Copyright
©
2017
Perry
Lee
Wood
Jr.

All
Rights
Reserved ABSTRACT

Phylogenetics,
Biogeography,
and
Patterns
of
Diversification
of
Geckos Across
the
Sunda
Shelf
with
an
Emphasis
on
the
Genus Cnemaspis
(Strauch,
1887)

Perry
Lee
Wood
Jr. Department
of
Biology,
BYU Doctor
of
Philosophy

In
my
dissertation
I
investigate
two
genera
of
geckos
(Cyrtodactylus
and
Cnemaspis)
that
are
distributed
across
Southeast
Asia
with
an
emphasis
on
Cnemaspis.
In
Chapter
1
I
use
a
multi- locus
dataset,
ancestral
area
analyses,
and
molecular
clock
dating
to
generate
a
species
level
time
calibrated
phylogeny
to
test
the
monophyly
of
Cyrtodactylus
and
to
identify
major
biogeographical
patterns.
I
identified
that
Cyrtodactylus
is
monophyletic
only
if
the
the
Sri
Lankan
genus
often
recognized
as
Geckoella
is
included.
The
results
of
the
Biogeographical
analyses
reveal
a
west
to
east
pattern
of
diversification.
Chapter
2
I
use
a
traditional
morphological
dataset
to
describe
a
new
species
of
Cnemaspis
from
Peninsular
Malaysia.
In
Chapter
3
my
colleagues
and
I
use
a
multi- locus
dataset
and
morphological
characters
to
revise
the
taxonomy
of
all
Cnemaspis
species
and
put
this
in
a
phylogenetic
context.
This
resulted
in
the
description
of
eight
new
species
and
allowed
us
to
generate
hypotheses
relating
to
parallel
evolution,
diversity,
and
biogeography.
In
Chapter
4
I
use
additional
taxon
sampling
of
Cnemaspis
from
Thailand
to
generate
a
more
complete
phylogeny
and
use
an
integrative
taxonomic
approach
to
describe
three
new
species.
In
Chapter
5
I
used
high
throughput
sequencing
of
sequence
capture
and
Ultra
Conserved
Elements
to
test
for
a
rapid
radiation
in
Cnemaspis
and
to
investigate
biogeographic
hypotheses
relating
to
the
diversification
and
evolution
of
Cnemaspis.
I
determined
that
there
was
a
temporal
rapid
radiation
of
Cnemaspis
that
coincides
with
the
temporal
diversification
of
other
terrestrial
vertebrates
across
Sundaland.
The
results
of
these
studies
indicate
that
the
species
diversity
of
Cnemaspis
is
underestimated
and
that
both
genera
have
experienced
similar
temporal
and
spatial
diversification
coinciding
with
major
vicariant
events
during
the
Eocene
and
the
Oligocene-Miocene
transition.

Keywords:
biogeography,
Cnemaspis,
Cyrtodactylus,
evolution,
Southeast
Asia,
taxonomy,
ultra
conserved
elements ACKNOWLEDGMENTS

I would like express many thanks to the graduate students at BYU, the department of biol- ogy (especially, Gentri Glaittli, Justina Tavana, and Dennis Shiozawa), and my committee (Byron

Adams, Seth Bybee, Duke Rogers, Lee Grismer, and Jack Sites) for helping me along the way. I would like to highlight a special thanks to my amazing and patient advisor, Jack Sites. He spent numerous hours and late nights/early mornings editing my manuscripts and making sure that I had everything I needed to excel and grow as a “great” graduate student at BYU. The funding for mul- tiple research trips and meetings was greatly appreciated, too. I am also grateful for my friend and colleague Lee Grismer who has been putting up with me for 16 years starting from a punk kid (un- dergraduate) to a graduate student and many years of field work together all over the world. I am grateful for the graduate students in the Site’s lab (César Aguilar, Derek Tucker, Luke Welton) and past students, post-doc Juan Santos and his wife Natalia Biani, and visiting researchers Mariana

Morando and Luciano Avila for many hours of conversations about research projects and normal day life. Thank you also to the people at the University of Washington (Adam Leaché, Jamie

Oaks, Leonard Jones, Itzue Caviedes-Solis, Nassima Bouzid, and Matt McElroy) for mentorship and emotional support while generating genomic data. I would like to acknowledge friends and colleagues Dan Portik and Phillip Skipwith, who provided advice and helpful discussions of phy- logenetic software. Special thanks to my international colleagues and friends, Evah Quah, Shahrul

Anuar, and Anchalee Aowphol (and their students) for sharing their love of science, great food, and hard work in the field. My family and friends for always being there for me regardless of the situation.

Lastly, my wife, Margaret, and my dog, Cloe, for putting up with me for leaving them all the time to go into the field multiple times a year. I could not do this without your unconditional love, support, and encouragement. Table of Contents

Title Page…………………………………………………………………………………………..i Abstract……………………………………………………………………………………………ii Acknowledgments………………………………………………………………………………..iii Table of Contents……………………………………………………………………………..iv–vii List of Tables………………………………………………………………………………...viii–ix List of Figures………………………………………………………………………………...x–xiv

Chapter 1: Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification…………………………………………………………………………………1–13 Abstract……………………………………………………………………………………2 Introduction……………………………………………………………………………..2–3 Materials and Methods………………………………………………………………….3–7 Taxon sampling, data collection, and sequence alignment………………………..3 Phylogeny reconstruction……………………………………………………….3–6 Divergence timing and ancestral area analyses………………………………...6–7 Results…………………………………………………………………………………..7–9 Discussion……………………………………………………………………………..9–12 Phylogeny……………………………………………………………………..9–10 Biogeography………………………………………………………………...10–11 Taxonomy……………………………………………………………………11–12 Conclusions………………………………………………………………………12 Acknowledgements………………………………………………………………………12 References……………………………………………………………………………12–13

Chapter 2: A new species of lowland karst dwelling Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from northwestern Peninsular Malaysia………………………………………14–35 Abstract…………………………………………………………………………………..15 Introduction………………………………………………………………………………15 Materials and Methods……………………………………………………………….17–18 Systematics…………………………………………………………………………..18–33 Discussion………………………………………………………………………………..33 Acknowledgments……………………………………………………………………….33 References……………………………………………………………………………33–34 Appendix……………………………………………………………………………..34–35

Chapter 3: Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia……………………………………………………………………………………36–183 Abstract…………………………………………………………………………...……...40 Introduction……………………………………………………………………...…..40–42 Materials and Methods……………………………………………………………….42–51 Molecular Analysis…………………………………………………………..42–43 Phylogenetic Analysis……………………………………………………………43 Morphological Analysis……………………………………………………...43–50

iv Species Accounts………………………………………………………………...51 Results……………………………………………………………………………....51–173 Taxonomy………………………………………………………………....……..58 Ca Mau Clade………………………………………………………………..58–62 Cnemaspis boulengerii………………………………………………59–60 Cnemaspis psychedelica……………………………………………..60–62 Pattani Clade…………………………………………………………………62–75 Cnemaspis monachorum………………………………………….….63–70 Cnemaspis biocellata………………………………………………...70–72 Cnemaspis niyomwanae………………………………………….…..72–74 Cnemaspis kumpoli…………………………………………………..74–75 Northern Sunda Clade..……………………………………………………..75–132 Chanthaburiensis Group…………………….……………………….76–95 Cnemaspis chanthaburiensis………...... ……………..76–78 Cnemaspis neangthyi……………………………………...... 78–79 Cnemaspis aurantiacopes…………………………………....79–80 Cnemaspis caudanivea……………………………………….80–82 Cnemaspis nuicamensis……………………………………...82–83 Cnemaspis tucdupensis……………………………………....84–85 Cnemaspis siamensis………………………………………...85–86 Cnemaspis huaseesom……………………………………….87–88 Cnemaspis chanardi………………………………………….88–90 Cnemaspis omari sp. nov…………………………………….90–94 Cnemaspis roticanai…………………………………………94–95 Argus Group………………………………………………………...96–102 Cnemaspis flavigaster………………………………………..96–97 Cnemaspis argus……………………………………………..98–99 Cnemaspis karsticola…………………………………………….99 Cnemaspis perhentianensis………………………………..100–102 Affinis Group………………………………………………………102–132 Cnemaspis affinis………………………………………….102–104 Cnemaspis harimau……………………………………….104–106 Cnemaspis pseudomcguirei……………………………….106–108 Cnemaspis shahruli………………………………………..108–110 Cnemaspis mcguirei……………………………………….110–114 Cnemaspis grismeri……………………………………….114–116 Cnemaspis flavolineata……………………………………116–117 Cnemaspis temiah sp. nov…………………………………117–121 Cnemaspis narathiwatensis……………………………….121–122 Cnemaspis hangus sp. nov………………………………...122–126 Cnemaspis selamatkanmerapoh………………………………...126 Cnemaspis bayuensis……………………………………...126–128 Cnemaspis stongensis sp. nov………………………..……129–132 Southern Sunda Clade……………………………………………………..133–161 Cnemaspis limi……………………………………………………133–134 Nigridia Group…………………………………………………….134–138

v Cnemaspis nigridia……………………………………………..136 Cnemaspis paripari………………………………………..136–138 Kendallii Group…………………………………………………...138–161 Cnemaspis sundainsula sp. nov…………………………...139–143 Cnemaspis kendallii……………………………………….143–147 Cnemaspis pemanggilensis………………………………..147–148 Cnemaspis baueri………………………………………….148–149 Cnemaspis mumpuniae sp. nov……………………………149–153 Cnemaspis peninsularis sp. nov…………………………...154–160 Cnemaspis bidongensis……………………………………160–161 Species Incertae Sedis…………………………………………………………..162 Cnemaspis laoensis………………………………………………..162–163 Cnemaspis punctatonuchalis…………………………………...... 163–164 Cnemaspis vandeventeri…………………………………………..165–166 Cnemaspis kamolnorranathi………………………………………166–167 Cnemaspis dringi………………………………………………….167–169 Cnemaspis sundagekko sp. nov……………………………………170–172 Discussion………………………………………………………………………....173–177 Comments on Biogeography……………………………………………....173–175 Comments on Diversity………………………………………………………...175 Comments on Parallel Evolution………………………………………….175–176 Comments on Genetic Divergence with Cnemaspis…………………………...177 Comments on Integrative Taxonomy…………………………………………...177 Acknowledgements……………………………………………………………………..177 References…………………………………………………………………………177–183

Chapter 4: Three new karst-dwelling Cnemaspis Strauch, 1887 (Squamata; Gekkonidae) from Peninsular Thailand and the phylogenetic placement of C. punctatonuchalis and C. vandeventeri……………………………………………………………………….184–230 Abstract……………………………………………………………………………185–186 Introduction………………………………………………………………………..186–187 Materials and Methods…………………………………………………………….189–191 Taxon Sampling………………………………………………………………...188 Molecular and Phylogenetic Analyses…………………………………….188–190 Morphological Analyses…………………………………………………..190–191 Results……………………………………………………………………………..191–222 Systematics………………………………………………………………..193–222 Cnemaspis lineogularis sp. nov…………………………………...193–205 Cnemaspis phangngaensis sp. nov………………………………..205–215 Cnemaspis thachanaensis sp. nov………………………………....215–222 Discussion…………………………………………………………………………222–223 Acknowledgments……………………………………………………………………....223 References…………………………………………………………………………225–230

Chapter 5: A Geologically-driven Rapid Radiation on the Sunda Shelf………………….231–293 Abstract…………………………………………………………………………………232

vi Introduction………………………………………………………………………..233–234 Results……………………………………………………………………………..234–238 Discussion…………………………………………………………………………238–240 Methods……………………………………………………………………………241–250 Data Collection……………………………………………………………241–242 Targeted Sequence Capture and UCE Data Collection…………………...242–243 Bioinformatics and Data Processing……………………………………....243–244 Phylogenomic Analyses…………………………………………………...244–247 Diversification and Ancestral Area Reconstructions……………………...247–249 References………………………………………………………………………....250–263 Acknowledgements……………………………………………………………………..264 Supplemental Material...…………………………...………………………………265–281

vii List of Tables Chapter 1 Table 1: Specimens used for phylogenetic analyses in this study…………....………...4–6 Table 2: Primers used for PCR amplification and sequencing……………………………7 Table 3: Best-fit models for data partitions as determined by AIC, and similar models chosen for phylogenetic analyses…………………………………………………………7

Chapter 2 Table 1: Diagnostic characters of Southeast Asian species of Cnemaspis…………..21–28 Table 2: Morphological data taken for the type series……………………....……….29–31

Chapter 3 Table 1: Specimens used for the molecular phylogenetic analyses...…………...…...45–50 Table 2: A list of primers used for amplification and sequencing……………………….51 Table 3: Selected and applied models of molecular evolution for data partitions determined by AIC for the phylogenetic analyses…………...…………………………..51 Table 4: Uncorrected p-distances for the genus Cnemaspis for the mitochondrial gene ND2 calculated in MEGAv5.2.2 (Tamura et al. 2011)………………………………56–57 Table 5: Uncorrected p-distances for the populations of Cnemaspis peninsularis sp. nov. for the mitochondrial gene ND2 calculated in MEGAv5.2.2 (Tamura et al. 2011)………………………………………………………………………………...…...64 Table 6: Diagnostic morphological characters separating various species of Cnemaspis from one another……………………………………….…………………….………65–67 Table 7: Diagnostic color pattern characters separating various species of Cnemaspis from one another………………………………………………..……………………68–69 Table 8: Meristic and mensural character states of the type series of Cnemaspis omari sp. nov……………………………………………………………………………………92–93 Table 9: Meristic and mensural character states of the type series of Cnemaspis temiah sp. nov……………………………………………….…………………………….119–120 Table 10: Meristic and mensural character states of the type series of Cnemaspis hangus sp. nov…………………………………………………………………..…………124–125 Table 11: Meristic and mensural character states of the type series of Cnemaspis stongensis sp. nov….……………………………………………………………...131–132 Table 12: Meristic and mensural character states of the type series of Cnemaspis sundainsula sp. nov………………………………………...…………………………...142 Table 13: Meristic and mensural character states of the type series of Cnemaspis mumpuniae sp. nov………………………………………………………...……...152–153 Table 14: Meristic and mensural character states of the type series of Cnemaspis peninsularis sp. nov……………………………………………..……………………...157 Table 15: Meristic and mensural character states of the type series of Cnemaspis sundagekko sp. nov………………….………………………………………………….171 Table 16: Activity period, substrate preference (microhabitat), elevation (upland > 600 m), and presence or absence of ocelli (eyespots) in the species of Cnemaspis for which we had sufficient data or observations...... 176

Chapter 4 Table 1: Models of molecular evolution used for the ML and BI analyses…………….190

viii Table 2: Pairwise uncorrected p-distances based on 1,335 bp of ND2 and associated tRNAs calculated in MEGA v6.06 (Tamura et al., 2013) within the chanthaburiensis group…………………………...……………………………………………………….193 Table 3: Pairwise uncorrected p-distances based on 1,335 bp of ND2 and associated tRNAs calculated in MEGA v6.06 (Tamura et al., 2013) within the siamensis group……………………….…………………………………………………………...193 Table 4: Mensural and meristic character states for the type series of Cnemaspis lineogularis sp. nov………………...……………………………………………...197–198 Table 5: Diagnostic color pattern characters separating various species of Cnemaspis from one another following Grismer et al., 2014d………………………………...199–201 Table 6: Diagnostic morphological characters separating C. lineogularis from species of Cnemaspis in the chanthaburiensis group………………………………………...202–203 Table 7: Mensural and meristic character states for the type series of Cnemaspis phangngaensis sp. nov…………………………………………………………….210–211 Table 8: Diagnostic morphological characters separating species of Cnemaspis from one another in the siamensis group…………………………………………………….212–213 Table 9: Menusural and meristic character state for the type series of Cnemaspis thachanaensis sp. nov……………………...……………………………………...218–219

Chapter 5 Table S1: Specimens used for the UCE/sequence capture dataset.……………….274–278 Table S2: The results of the model fitting from the ancestral area reconstructions in BioGeoBears. d= is the rate of dispersal or range expansion, e= is the extinction rate, and j=is the jump-dispersal rate……………………………………………………………..279

ix List of Figures Chapter 1 Figure 1: Global distribution of Cyrtodactylus, with place names mentioned in the text listed……………………………………………………………………………………….3 Figure 2: Maximum likelihood phylogeny of Cyrtodactylus, under the 13-partition scheme (ln L 71587.112627)…..………………………………………………………….8 Figure 3: Historical biogeography of Cyrtodactylus……….……………………………10

Chapter 2 Figure 1: Distribution map showing the location of the type locality of Cnemaspis grismeri sp. nov. in the Lenggong Valley, Perak………………………………………..16 Figure 2: Upper left: adult female Cnemaspis grismeri sp. nov. paratype LSUHC 9973 in dark daytime coloration.………………………………………………………………....19 Figure 3: Differences between male and female Cnemaspis grismeri sp. nov. in the development of the dark shoulder patch…………………………………………………20 Figure 4: Microhabitat structure of the karst formations for Cnemaspis grismeri sp. nov. from Lenggong Valley, Perak……………………………………………………………32

Chapter 3 Figure 1: Distribution of the Sundaland Rock Gecko genus Cnemaspis Strauch, 1887 in Southeast Asia……………………………………………………………………………41 Figure 2: Maximum-likelihood phylogram (–InL 73957.608688) of the species of the genus Cnemaspis with Bayesian posterior probabilities (BPP) and maximum-likelihood (ML) bootstrap values, respectively based on the concatenated 3 gene dataset…………52 Figure 3: Distribution of the species and species groups in the Ca Mau and Northern Sunda clades……………………………...………………………………………………53 Figure 4: Distribution of the species in the Pattani and Southern Sunda clades…………54 Figure 5. Ecological parameters of activity period, elevation (upland species are > 600 m in elevation), substrate preference (microhabitat), and the presence or absence of ocelli (eyespots) mapped onto the molecular phylogeny (Fig. 2) of the species of Cnemaspis………………………………………………………………………………..55 Figure 6: Adult male Cnemaspis boulengerii (LSUDPC 8199) from Con Son Island in the Con Dao Archipelago, Ba-Ria-Vung Tau Province, Vietnam in the dark color pattern phase……………………..………………………………………………………………58 Figure 7: Cnemaspis boulengerii from Con Son Island in the Con Dao Archipelago, Ba- Ria-Vung Tau Province, Vietnam……………………………………………………..…59 Figure 8: Cnemaspis psychedelica from Hon Khoai Island, Ca Mau Province, Vietnam………………………………………………….……………………………….61 Figure 9: Adult male Cnemaspis monachorum (LSUDPC 6502) from Pulau Langgun of the Langkawi Archipelago, Kedah, Peninsular Malaysia in the dark color pattern phase……………...……………………………………………………………………...62 Figure 10: Cnemaspis monachorum (LSUDPC 6502) from the Langkawi Archipelago, Kedah, Peninsular Malaysia….…………………………………………………………..63 Figure 11: Cnemaspis biocellata from Perlis, Peninsular Malaysia.…………………….71 Figure 12: Cnemaspis biocellata from Perlis, Peninsular Malaysia.…………………….72 Figure 13: Cnemaspis niyomwanae from Thum Khao Ting, Palean District, Trang

x Province, Thailand…………………...…………………………………………………..73 Figure 14: Cnemaspis kumpoli….………………………………………………………..75 Figure 15: Cnemaspis chanthaburiensis from the Phnom Samkos Wildlife Sanctuary, Pursat Province, Cardamom Mountains, Cambodia…...………………………………...76 Figure 16: Habitat of Cnemaspis chanthaburiensis at Phnom Dalai, Phnom Samkos Wildlife Sanctuary, Pursat Province, Cardamom Mountains, Cambodia...... …………77 Figure 17. Upper left: adult male Cnemaspis neangthyi from near the village of O’lakmeas, Pursat Province, Cambodia………………………………...………………..78 Figure 18. Cnemaspis aurantiacopes from Hon Dat Hill, Kien Giang Province, Vietnam...…………...……………………………………………………………………80 Figure 19. Cnemaspis caudanivea from Hon Tre Island, Kien Giang Province, Vietnam……………………………………………… ………………………………….81 Figure 20. Upper; adult male (LSUDPC 3136) in the light color pattern phase….……..83 Figure 21. Cnemaspis tucdupensis from Tuc Dup Hill, An Giang Province, Vietnam...... 84 Figure 22. Cnemaspis siamensis from Pathio, Chumpon Province, Thailand…………...86 Figure 23. Cnemaspis huaseesom from Sai Yok National Park, Kanchanaburi Province, Thailand……………………………………………….…………………………………87 Figure 24. Left: karst microhabitat of Cnemaspis huaseesom at Sai Yok National Park, Kanchanaburi Province, Thailand...……………………………………………………..88 Figure 25. Left: tree hole microhabitat of Cnmeaspis chanardi at the Phuphaphet Cave area, Satun Province, Thailand..…………………………………………………………89 Figure 26. Cnemaspis omari sp. nov…...………………………………………………..91 Figure 27. Lowland dipterocarp forest of Cnemaspis omari sp. nov. at Perlis State Park, Perlis, Peninsular Malaysia…………………….………………………………………...94 Figure 28. Cnemaspis roticanai from Gunung Raya, Pulau Langkawi, Kedah, Peninsular Malaysia…………………………………………………………………………..……...95 Figure 29. Cnemaspis flavigaster from FRIM, Kepong, Selangor, Peninsular Malaysia……………………………………………………..…………………………...97 Figure 30. Left: microhabitat of Cnemaspis argus on Gunung Lawit, Terengganu, Peninsular Malaysia……….……………………………………………………………..98 Figure 31. Cnemaspis karsticola from Gunung Reng, Kelantan, Peninsular Malaysia………………………………………………………………………………….99 Figure 32. Upper: adult male Cnemaspis perhentianensis (LSUDPC 5159) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia in the dark color pattern phase…………………….……………………………………………………………...101 Figure 33. Cnemaspis affinis from Penang Hill, Pulau Pinang, Penang, Peninsular Malaysia………………………………………………………………………………...103 Figure 34. Cnemaspis harimau from Sungai Badak, Gunung Jerai, Kedah, Peninsular Malaysia.………………………………………………………………………………..104 Figure 35. Microhabitat of Cnemaspis harimau at Sungai Badak, Gunung Jerai, Kedah, Peninsular Malaysia……………………..……………………………………………...105 Figure 36. Cnemaspis pseudomcguirei from Bukit Larut, Perak, Peninsular Malaysia………………………………………………………………………………...107 Figure 37. Microhabitat of Cnemaspis pseudomcguirei at Bukit Larut, Perak, Peninsular Malaysia……………………..………………………………………………………….108

xi Figure 38. Cnemaspis shahruli from Peninsular Malaysia..……………………………109 Figure 39. Adult male Cnemaspis mcguirei (LSUDPC 5173) from Bukit Larut, Perak, Peninsular Malaysia in the light color pattern phase……………………..…………….110 Figure 40. Cnemaspis mcguirei (LSUDPC 5173) from Bukit Larut, Perak, Peninsular Malaysia………………………….……………………………………………………..111 Figure 41. Cnemaspis grismeri from Gua Asar, Bukit Kepala Gajah, Lenggong, Perak, Malaysia………………………………………………………………………………...112 Figure 42. Upper: juvenile Cnemaspis grismeri (LSUDPC 6695) from Gua Asar, Bukit Kepala Gajah limestone massif, Lenggong, Perak, Malaysia in light color pattern phase……………………………………………………………...…………………….113 Figure 43. Upper: adult male Cnemaspis flavolineata (LSUHC 8079) from the Gap at Fraser’s Hill, Pahang, Peninsular Malaysia.……………………………………………115 Figure 44. Cnemaspis temiah sp. nov. from Tanah Rata, Cameron Highlands, Pahang, Peninsular Malaysia…………...………………………………………………………..118 Figure 45. Cnemaspis narathiwatensis from Sungai Enam, Belum, Perak, Peninsular Malaysia...…………………………………………...………………………………….122 Figure 46. Cnemaspis hangus sp. nov. from Bukit Hangus, Pahang, Peninsular Malaysia…………………………………………………..…………………………….123 Figure 47. Upper: adult male holotype (LSUHC 11016) Cnemaspis selamatkanmerapoh sp. nov. in the dark color pattern phase………………………………………………...127 Figure 48. Cnemaspis bayuensis from Gua Bayu, Kelantan, Peninsular Malaysia………………………………………………………………………………...128 Figure 49. Cnemaspis stongensis sp. nov. from Kem Baha, Gunung Stong, Peninsular Malaysia…………...……………………………………………………………………130 Figure 50. Cnemaspis limi from Pulau Tioman, Pahang, Peninsular Malaysia.………..134 Figure 51. Cnemaspis nigridia from Gunung Gading, Sarawak, East Malaysia..……..135 Figure 52. Adult male Cnemaspis paripari (LSUDPC 4920) from Gua Pari-pari, Sarawak, East Malaysia in the dark color pattern phase……………………...………...137 Figure 53. Cnemaspis paripari from Gua Angin, Sarawak, East Malaysia...…………138 Figure 54. Cnemaspis sundainsula sp. nov. from Mount Ranai, Kecamatan Bunguran Timur, Kabupaten Natuna, Propinsi Kepulauan Riau, Bunguran (Great Natuna) Island, Indonesia...... …………………...140 Figure 55. Cnemaspis kendallii………………………..………….……………………144 Figure 56. Upper: adult male Cnemaspis kendallii (LSUDPC 4901) at Santubong, Sarawak, East Malaysia in the dark color pattern phase displaying the white underside of the original tail………………………………………………………………………….145 Figure 57. Upper: adult male Cnemaspis pemanggilensis (LSUDPC 2578) from Pulau Pemanggil, Johor, Peninsular Malaysia in the dark color pattern phase………………………………….………………………………………………...146 Figure 58. Cnemaspis baueri from Pulau Aur, Johor, Peninsular Malaysia…………..148 Figure 59. Cnemaspis mumpuniae sp. nov. from Pulau Natuna Besar…………………150 Figure 60. Cnemaspis peninsularis sp. nov. from Gunung Ledang, Johor, Peninsular Malaysia……...………………………………………………………………………....154 Figure 61. Cnemaspis peninsularis sp. nov. …..………………………………...…….155

xii Figure 62. Maximum likelihood phylogram (–InL 73957.608688) of Cnemaspis peninsularis sp. nov. based on the mitochondrial gene ND2 and the geographic distribution of the specimens sampled……………... ………………………………….158 Figure 63. Cnemaspis bidongensis from Pulau Bidong, Terengganu, Peninsular Malaysia…………………………………………………...……………………………161 Figure 64. Adult female holotype of Cnemaspis laoensis (THNHM 12433) from Dong Phu Vieng National Protected Area, Savannakhet, Laos……………………………….162 Figure 65. Cnemaspis punctatonuchalis from Thap Sakae District, Prachuap Khirikhan Province, Thailand……………………………………………………………………...164 Figure 66. Cnemaspis vandeventeri from Phuket Island, Phuket Province, Thailand…………………...……………………………………………………………165 Figure 67. Cnemaspis kamolnorranthi from the Petchphanomwat Waterfall, in Tai Rom Yen National Park, Surat Thani Province, Thailand……………………………………166 Figure 68. Type series of Cnemaspis dringi from Sarawak, East Malaysia...………….168 Figure 69. Type series of Cnemaspis sundagekko sp. nov. from Pulau Siantan, Anambas Archipelago, Riau Province, Indonesia…………………..…………………………….169

Chapter 4 Figure 1: Distribution of the species of Cnemaspis in the chanthaburiensis and siamensis groups………………………………...…………………………………………………189 Figure 2: Phylogenetic relationships of the chanthaburiensis (A) and the siamensis (B) groups………………………………...…………………………………………………192 Figure 3: (A) male holotype BYU 62535 and (B) female paratype ZMKU R 00728 of Cnemaspis lineogularis sp. nov………………………………………………………...194 Figure 4: Ventral coloration and sexual dichromatism in the type series of Cnemaspis lineogularis sp. nov……………………………….…………………………………….195 Figure 5: Dorsal view of the type series of Cnemaspis lineogularis sp. nov...…………196 Figure 6: (A) habitat and (B) microhabitat of Cnemaspis lineogularis sp. nov………..204 Figure 7: (A) adult male holotype BYU 62538 and (B) female paratype BYU 62537 of Cnemaspis phangngaensis sp. nov……………………………………………………..206 Figure 8: Ventral coloration and sexual dichromatism of Cnemaspis phangngaensis sp. nov………………………………………………………………………………………207 Figure 9: Dorsal coloration of the type series of Cnemaspis phangngaensis sp. nov………………………………………………………………..……………………..208 Figure 10: (A) general karst and limestone forest near the type locality of Cnemaspis phangngaensis sp. nov. (B) karst microhabitat where C. phangngaensis occurs. …….214 Figure 11: Coloration of Cnemaspis thachanaensis sp. nov…..……………………….216 Figure 12: Ventral coloration and sexual dichromatism of the type series of Cnemaspis thachanaensis sp. nov., males: (A) BYU 62543, (B) BYU 62544 (holotype), (C) ZMKU R 00731, females: (D) ZMKU R 00729, (E) ZMKU R 00730, (F) BYU 62542………217 Figure 13: Dorsal coloration of the type series of Cnemaspis thachanaensis sp. nov., males: (A) BYU 62543, (B) BYU 62544 (holotype), (C) ZMKU R 00731, females: (D) ZMKU R 00729, (E) ZMKU R 00730, (F) BYU 62542……………………………….217 Figure 14: (A) karst and limestone forest near the type locality of Cnemaspis thachanaensis sp. nov…………..………………………………………………………220

xiii Chapter 5 Figure 1: A. Chronogram of Cnemaspis based on a concatenated ExaBayes analysis of 514 loci guide tree (full tree in supplemental Figure S5)…..…………………………..271 Figure S1: Distribution of the genus Cnemaspis based on Grismer et al. 2014; colors identify major clades…...……………………………………………………………….272 Figure S2: Maximum Likelihood trees estimated in IQTREE (Nguyen et al. 2015; Minh et al. 2013) using three different sized datasets: A. 50p (267,912 bp/514 loci), B. 60p (231,027 bp/440 loci), C.75p (57,940bp/105 loci)……………………………………..273 Figure S3: Phylogenomic relationships of Cnemaspis based on 514 loci (267,912 bp)…………………………………………………………………...... ………………..274 Figure S4: SVDQuartets consensus species tree with bootstrap consensus support values (SVDBS)………………………………………………………………………………..275 Figure S5: Numbers near the nodes are mean divergence time estimates from fossil and root height calibrations implemented in MCMCtree………………….………………..276 Figure S6: Estimated net speciation and extinction rates for Cnemaspis from BAMM outputs based on the time-calibrated tree..……………………………………………...277 Figure S7: The BAMM tree with a single rate shift (blue circle, probability=0.70) associated with a rapid radiation of Cnemaspis with the exclusion of the Ca Mau clade…………………………………………………………………………………….278 Figure S8: A. Lineage through time plot based on the time-calibrated tree generated from MCMCtree...……………………………………………………………………………279 Figure S9: Summary statistics from the raw Illumina reads...………………………….280

xiv Chapter 1: Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of

diversification

1 Molecular Phylogenetics and Evolution 65 (2012) 992–1003

Contents lists available at SciVerse ScienceDirect

Molecular Phylogenetics and Evolution

journal homepage: www.elsevier.com/locate/ympev

Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification

1 Perry L. Wood Jr. , Matthew P. Heinicke ⇑, Todd R. Jackman, Aaron M. Bauer

Department of Biology, 147 Mendel Hall, Villanova University, 800 Lancaster Ave., Villanova, PA 19085, USA article info abstract

Article history: The Asian/Pacific genus Cyrtodactylus is the most diverse and among the most widely distributed genera Received 7 May 2012 of geckos, and more species are continually being discovered. Major patterns in the evolutionary history Revised 24 August 2012 of Cyrtodactylus have remained largely unknown because no published study has broadly sampled across Accepted 27 August 2012 the geographic range and morphological diversity of the genus. We assembled a data set including Available online 13 September 2012 sequences from one mitochondrial and three nuclear loci for 68 Cyrtodactylus and 20 other gekkotan spe- cies to infer phylogenetic relationships within the genus and identify major biogeographic patterns. Our Keywords: results indicate that Cyrtodactylus is monophyletic, but only if the Indian/Sri Lankan species sometimes Sundaland recognized as Geckoella are included. Basal divergences divide Cyrtodactylus into three well-supported Indonesia Biogeography groups: the single species C. tibetanus, a clade of Myanmar/southern Himalayan species, and a large clade Dispersal including all other Cyrtodactylus plus Geckoella. Within the largest major clade are several well-supported Myanmar subclades, with separate subclades being most diverse in Thailand, Eastern Indochina, the Sunda region, Papuan region the Papuan region, and the Philippines, respectively. The phylogenetic results, along with molecular clock and ancestral area analyses, show Cyrtodactylus to have originated in the circum-Himalayan region just after the Cretaceous/Paleogene boundary, with a generally west to east pattern of colonization and diver- sification progressing through the Cenozoic. Wallacean species are derived from within a Sundaland radi- ation, the Philippines were colonized from Borneo, and Australia was colonized twice, once via New Guinea and once via the Lesser Sundas. Overall, these results are consistent with past suggestions of a Palearctic origin for Cyrtodactylus, and highlight the key role of geography in diversification of the genus. Ó 2012 Elsevier Inc. All rights reserved.

1. Introduction 2009, 2010a, 2010b); and Wallacea, Australia, and New Guinea (Kraus, 2008; Oliver et al., 2011; Shea et al., 2012). Cyrtodactylus Gray, 1827 (bent-toed geckos), with more than Despite great activity in terms of alpha systematic studies, phy- 150 recognized species, is by far the most species-rich genus of logenetic research on Cyrtodactylus has been relatively limited thus gekkotan lizards (Uetz, 2012). Recently, as many as 19 new species far. Regional phylogenies with reasonably broad taxon sampling have been described in a given year from throughout the group’s have been generated for the bent-toed geckos of the Philippines broad range in Asia and the western Pacific (Fig. 1), and since the (Siler et al., 2010; Welton et al., 2010a, 2010b), Australia and Mel- start of the 21st century, known diversity of Cyrtodactylus has more anesia (Shea et al., 2012), and for some Malay Peninsula and than doubled. Virtually all regions occupied by Cyrtodactylus have Sundaland Cyrtodactylus (Grismer et al., 2010). Monophyly and seen a huge increase in the number of recognized species de- interrelationships of the bent-toed geckos of these and other geo- scribed, including Myanmar (Bauer, 2002, 2003; Mahony, 2009), graphic regions has yet to be established. Indeed, the composition Vietnam, (Ngo, 2008; Ngo and Bauer, 2008; Nguyen et al., 2006), of Cyrtodactylus as a whole remains unclear, especially with respect Sundaland (Chan and Norhayati, 2010; Grismer et al., 2010; Iskan- to certain taxa in Nepal, northern India, and Tibet, which have been dar et al., 2011; Oliver et al., 2009), the Philippines (Welton et al., variously assigned to Cyrtodactylus, Cyrtopodion, Altigekko, Altiphy- lax, Indogekko, and Siwaligekko (see Krysko et al. (2007) for a recent Geckoella Corresponding author. Present address: Department of Natural Sciences, review). In addition, the status of , a presumably ⇑ University of Michigan-Dearborn, 125 Science Building, 4901 Evergreen Road, monophyletic group of small, ground-dwelling bent-toed geckos Dearborn, Michigan 48128, USA. (regarded as either a genus or a subgenus of Cyrtodactylus; Kluge, E-mail addresses: [email protected] (P.L. Wood Jr.), heinicke@umd. 2001; Bauer, 2002) remains uncertain. umich.edu (M.P. Heinicke), [email protected] (T.R. Jackman), aaron. A practical problem engendered by the lack of broader scale [email protected] (A.M. Bauer). 1 phylogenetic resolution in Cyrtodactylus is that each newly de- Present address: Department of Biology, Brigham Young University, Provo, UT 84602, USA. scribed species must be diagnosed relative to all of its congeners,

1055-7903/$ - see front matter Ó 2012 Elsevier Inc. All rights reserved. http://dx.doi.org/10.1016/j.ympev.2012.08.025 2 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003 993

Fig. 1. Global distribution of Cyrtodactylus, with place names mentioned in the text listed. or the assumption must be made that geographically coherent and themselves or photographic vouchers were examined by one or morphologically similar species are monophyletic. While this ap- more authors but in some cases we were dependent on the species pears to be the case in some instances, it is likely that not all such identifications of collectors or other institutions. Given the rapid groups are natural, and without broad sampling monophyly cannot rate of description of Cyrtodactylus spp. and the break-up of ‘‘spe- be conclusively demonstrated even for regional clades that do ap- cies’’ previously believed to be widespread (e.g. Johnson et al., pear monophyletic (e.g., Philippines: Siler et al., 2010; Welton 2012), it is possible that some identifications may need subsequent et al., 2010a, 2010b; Queensland: Shea et al., 2012). A phylogeny revision, however, all may be considered accurate to at least spe- spanning the geographic distribution and morphological variation cies group. Genomic DNA was extracted from tissue samples using of Cyrtodactylus and having sufficient sampling to provide a ‘‘back- Qiagen DNeasy™ tissue kits under manufacturers’ protocols. All bone’’ for the genus would make a major contribution in providing genes were amplified using a double-stranded Polymerase Chain a preliminary estimate of the monophyly of as yet unevaluated Reaction (PCR). Included in the reaction were 2.5 ll genomic presumptive clades. In addition, such a phylogeny can provide DNA, 2.5 ll light strand primer 2.5 ll heavy strand primer, 2.5 ll information about relevant outgroups for future regional Cyrto- dinucleotide pairs, 2.5 ll5 buffer, MgCl 10 buffer, 0.18 ll Taq Ó Ó dactylus phylogenies, allow evaluation of the generic allocation of polymerase, and 9.82 llH2O, using primers listed in Table 2. PCR the problematic taxa that have been variously assigned to Cyrto- reactions were executed on an Eppendorf Mastercycler gradient dactylus or other genera, and provide a framework for comparative theromocycler under the following conditions: initial denaturation analyses of Cyrtodactylus biology, including biogeography and mor- at 95 °C for 2 min, followed by a second denaturation at 95 °C for phological evolution. To this end we used nucleotide sequence data 35 s, annealing at 50–55 °C for 35 s, followed by a cycle extension from approximately 45% of recognized bent-toed gecko species, at 72 °C for 35 s, for 34 cycles. All PCR products were visualized via including exemplars encompassing the morphological range of 1.5% agarose gel electrophoresis. Successful PCR amplifications variation and from across the geographic range of Cyrtodactylus, were purified using AMPure magnetic bead solution (Agencourt to erect such a backbone phylogeny for the genus. We use this data Bioscience). Purified PCR products were sequenced using Applied set to evaluate current taxonomy and make preliminary observa- Biosystems BigDye™ Terminator v3.1 Cycle Sequencing ready tions of Cyrtodactylus historical biogeography. reaction kit or DYEnamic™ ET Dye Terminator kit (GE Healthcare). Products were purified using a Cleanseq magnetic bead solution (Agentcourt Bioscience). Purified sequence reactions were ana- 2. Materials and methods lyzed using an ABI 3700 or ABI 3730XL automated sequencer. All sequences were analyzed from the 3 and the 5 ends indepen- 2.1. Taxon sampling, data collection, and sequence alignment 0 0 dently to ensure congruence between the reads. The forward and the reverse sequences were imported and edited in Geneious™ We constructed a nucleotide sequence dataset for 68 species of version v5.4 (Drummond et al., 2011); ambiguous bases were Cyrtodactylus from throughout the range of the genus, plus two corrected by eye. All edited sequences were aligned by eye. Pro- species of Geckoella, six Hemidactylus, and six genera and species tein-coding sequences were investigated in MacClade v4.08 of Palearctic naked-toed gecko. For purposes of establishing a time- (Maddison and Maddison, 2003) to ensure the lack of premature tree for the group, we included six additional gekkotan outgroups stop codons and to calculate the correct amino acid reading frame. as well as Anolis and Python (Table 1). Three outgroups – Python, Lialis, and Stenodactylus – are composites of two closely related species. The dataset consists of the complete mitochondrial gene 2.2. Phylogeny reconstruction ND2 and flanking tRNAs (Ala, Asn, Cys, Tyr), plus portions of the nuclear genes RAG1, PDC, and MXRA5. New sequences are depos- For comparative purposes, phylogenetic reconstructions were ited under GenBank accession numbers JX440515–JX440726. implemented using one character-based approach, Maximum Par- Liver, muscle, or tail tissue samples were derived from individ- simony (MP) and two model-based approaches, Maximum Likeli- uals collected in the field by the authors or donated by other hood (ML) and Bayesian Inference (BI). Maximum Parsimony researchers (see acknowledgments). When possible, specimens (MP) phylogeny and bootstrap estimates for nodal support were 3 Table 1 994 Specimens used for phylogenetic analyses in this study. Identification numbers are abbreviated as follows: ACD, Arvin C. Diesmos field collection; AdS, Anslem de Silva field series (specimens pending accession at the National Museum of Sri Lanka); AMS, Australian Museum, Sydney; BPBM, Bernice P. Bishop Museum; CAS, California Academy of Sciences; CES, Center for Ecological Sciences, Indian Institute of Sciences, Bangalore; CJS, Christopher J. Schneider field series; CUMZ, Chulalongkorn University Museum of Zoology; FK, Fred Kraus field series; FMNH, Field Museum of Natural History; ID, Indraneil Das field series; IRSNB, Institute des Sciences Naturelles du Belgique, Brussels; JB, Jon Boone captive collection; JFBM, James Ford Bell Museum of Natural History (Minnesota); KU, Kansas University Museum of Natural History; LSUHC, La Sierra University Herpetological Collection; LSUMZ, Louisiana State University Museum of Zoology; MAM, Mohammed Al-Mutairi field series; MFA, M. Firoz Ahmed field series; MVZ, Museum of Vertebrate Zoology (Berkeley); RAH, Rod A. Hitchmough field series; RMBR, Rafe M. Brown field series; SP, Sabah Parks Reference Collection; TNHC, Texas Natural History Collection; USNM, United States National Museum (Smithsonian); WAM, Western Australian Museum; YPM, Yale Peabody Museum; ZRC, Zoological Reference Collection, Raffles Museum.

ID number Species Locality GenBank accession numbers ND2 MXRA5 PDC RAG1 n/a Anolis carolinensis n/a EU747728 AAWZ 02008741 AAWZ 02013979 AAWZ 02015549 n/a Python molurus n/a AEQU 010243110 AEQU 01027927 AEQU 010344888 n/a Python regius n/a AB177878 FMNH 247474 Agamura persica Pakistan, Balochistan, Makran district, Gwadar division JX440515 JX440566 JX440625 JX440675 CAS 228737 Bunopus tuberculatus United Arab Emirates, Sharjah JQ945355 JX440676 MVZ 234350 Bunopus tuberculatus Iran, Qeshm Island HQ443540 JB 127 Cyrtopodion elongatum captive JX440516 JX440567 JX440626 JX440677 992–1003 (2012) 65 Evolution and Phylogenetics Molecular / al. et Jr. Wood P.L. CES 08022 Hemidactylus anamallensis India, Tamil Nadu, Ervikulam HM662368 HM622353 MVZ 245438 Hemidactylus angulatus Ghana, Volta region, Togo Hills EU268367 EU268336 EU268306 CAS 229633 Hemidactylus frenatus Myanmar, Tanintharyi Division, Kaw Thaung District HM559629 HM559662 HM559695 AMS R167808 Hemidactylus frenatus New Caledonia, Sommet Poum JX440568 CAS 223286 Hemidactylus garnotii Myanmar, Rakhine State, Taung Gok Township EU268363 EU268332 EU268302 AMS R167800 Hemidactylus garnotii New Caledonia, Sommet Poum JX440569 YPM 14798 Hemidactylus mabouia USA, Florida HM559639 HM559672 HM559705 LSUMZ H-1981 Hemidactylus turcicus USA, Louisiana EU268360 EU268329 EU268299 AMS 141027 Lialis burtonis Australia, Victoria, Beulah Station JX440570 JFBM 8 Lialis burtonis Australia GU459742 GU459540 n/a Lialis jicari Australia AY369025 n/a Mediodactylus russowii captive JX440517 JX440627 JX440678 AMS 143861 Oedura marmorata Australia, Queensland GU459951 JX440571 EF534819 EF534779 MVZ 197233 Pygopus nigriceps Australia, Northern Territory JX440518 JX440572 EF534823 EF534783 CAS 198428 Sphaerodactylus roosevelti USA, Puerto Rico JN393943 JX440573 EF534825 EF534785 JB 34 Sphaerodactylus torrei Cuba JX440519 JX440574 EF534829 EF534788 JB 35 Stenodactylus petrii captive JX440628 JX440679 MVZ 238919 Stenodactylus petrii Niger, 49 km S Agadez HQ443548 MAM 3066 Stenodactylus slevini Saudi Arabia, Ibex Reserve JX440575 JB 28 Tropiocolotes steudneri captive JX440520 JX440576 JX440629 JX440680 RAH 292 Woodworthia maculata New Zealand, Titahi Bay GU459852 JX440577 GU459651 GU459449 JB 7 Cyrtodactylus (Geckoella) deccanensis captive (from Indian stock) JX440521 JX440630 JX440681 AdS 35 Cyrtodactylus (Geckoella) triedra Sri Lanka, Yakkunehela JX440522 JX440578 JX440631 JX440682 n/a Cyrtodactylus adorus Australia, Northeast Queensland HQ401166 KU 310100 Cyrtodactylus agusanensis Philippines, Dinagat Island, Municipality of Loreto GU550818 FMNH 265815 Cyrtodactylus angularis Thailand, Sa Kaeo, Muang Sa Kaeo JX440523 JX440579 JX440632 JQ945301 CAS 215722 Cyrtodactylus annandalei Myanmar, Sagging Division, Alaung Daw Kathapa National JX440524 JX440633 JX440683 Park, Gon Nyin Bin Camp ACD 2637 Cyrtodactylus annulatus Philippines, Mindanao Island, Eastern Mindanao, Diwata GU366085 Mountain Range LSUHC 7286 Cyrtodactylus aurensis West Malaysia, Johor, Pulau Aur, behind kg. Berhala JX440525 JX440580 JX440684 CAS 216459 Cyrtodactylus ayeyarwadyensis Myanmar, Rakhine State, Than Dawe District, Gwa Township, JX440526 JX440581 JX440634 JX440685 Rakhine Yoma Elephant Range, Elephant Camp SP 06906 Cyrtodactylus baluensis Malaysia, Borneo, Sabah, Mt. Kinabalu National Park, GU366080 Headquarters LSUHC 8933 Cyrtodactylus batucolus West Malaysia, Melaka, Pulau Besar JQ889178 JX440582 JX440635 JX440686 CAS 214104 Cyrtodactylus brevidactylus Myanmar, Mandalay Division, Popa Mountain Park, Kyauk JX440527 JX440583 JX440636 JX440687 Pan Tawn Township LSUHC 4056 Cyrtodactylus cavernicolus East Malaysia, Sarawak, Niah Cave JX440528 CUMZ 2003.62 Cyrtodactylus chanhomeae Thailand, Saraburi Province, Phraputthabata District, Khun JX440529 JX440584 JX440637 JX440688 Khlon Subdistrict, Thep Nimit Cave 4 CAS 226141 Cyrtodactylus chrysopylos Myanmar, Shan State, Ywa Ngan Township, Panlaung- JX440530 JX440585 JX440638 JX440689 Pyadalin Cave Wildlife Sanctuary LSUHC 8595 Cyrtodactylus cf. condorensis Vietnam, Kien Giang Province, Kien Hai District, Hon Son JX440531 JX440586 JX440639 JX440690 Island LSUHC 6546 Cyrtodactylus consobrinus West Malaysia, Selangor, Kepong, FRIM JX440532 JX440587 JX440640 JX440691 WAM R98393 Cyrtodactylus darmandvillei Indonesia, Gua, 7 km NW Sumbawa Besar JX440533 JX440588 JX440641 JX440692 LSUHC 8598 Cyrtodactylus eisenmanae Vietnam, Kien Giang Province, Kien Hai District, Hon Son JX440534 JX440589 JX440642 JX440693 Island LSUHC 6471 Cyrtodactylus elok West Malaysia, Pahang, Fraser’s Hill, the Gap JQ889180 JX440590 JX440643 JX440694 BPBM 18654 Cyrtodactylus epiroticus Papua New Guinea, Morobe Province, Apele, Mt. Shungol JX440535 JX440591 JX440644 JX440695 CES 091196 Cyrtodactylus fasciolatus India, Uttarkhand, Mussoorie HM622366 HM622351 USNM 559805 Cyrtodactylus feae Myanmar,Mandalay Division, Popa (village), vicinity of Popa JX440536 JX440592 JX440645 JX440696 Mountain Park CAS 222412 Cyrtodactylus gansi Myanmar, Chin State, Min Dat District, Min Dat Township, JX440537 JX440593 JX440646 JX440697 Che stream LSUHC 8638 Cyrtodactylus grismeri Vietnam, An Giang Province, Tuc Dup Hill JX440538 JX440594 JX440647 JX440698

LSUHC 8583 Cyrtodactylus hontreensis Vietnam, Kien Giang Province, Kien Hai District, Hon Tre JX440539 JX440595 JX440699 992–1003 (2012) 65 Evolution and Phylogenetics Molecular / al. et Jr. Wood P.L. Island n/a Cyrtodactylus hoskini Australia, Northeast Queensland HQ401119 FMNH 255454 Cyrtodactylus interdigitalis Lao PDR, Khammouan Province, Nakai District JQ889181 JX440596 JX440648 JX440700 FMNH 265812 Cyrtodactylus intermedius Thailand, Sa Kaeo, Muang Sa Kaeo JQ889182 JX440597 JX440649 JX440701 FMNH 258697 Cyrtodactylus irregularis Lao PDR, Champasak Province, Pakxong District JX440540 JX440598 JX440650 JQ945302 KU 314793 Cyrtodactylus jambangan Philippines, Mindanao Island, Zamboanga del Sur Prov., GU366100 Municipality of Pasonanca, Pasonanca Natural Park, Tumaga River FMNH 255472 Cyrtodactylus jarujini Lao PDR, Bolikhamxay Province, Thaphabat District JX440541 JX440599 JX440651 JQ945303 MVZ 239337 Cyrtodactylus jellesmae Indonesia, Sulawesi Island, Propinsi Sulawesi Selatan, JX440542 JX440600 JX440652 JX440702 Kabupaten Luwu Utara, Kecematan Malili, ca. 4 km N of Malili MFA 50083 Cyrtodactylus khasiensis India, Assam, Kaziranga, Kohora, Haldhibari JX440543 WAM R164144 Cyrtodactylus kimberleyensis Australia, Western Australia, East Montalivet Island JX440544 JX440601 JX440653 JX440703 n/a Cyrtodactylus klugei Papua New Guinea, Sudest Island HQ401198 FK 7709 Cyrtodactylus loriae Papua New Guinea, Milne Bay Prov., Bunisi, N slope Mt. EU268350 JX440602 EU268319 EU268289 Simpson n/a Cyrtodactylus louisiadensis Papua New Guinea, Sudest Island HQ401190 LSUHC 7532 Cyrtodactylus macrotuberculatus Malaysia, Kedah, Pulau Langkawi, Gunung Raya JX440545 JX440603 JX440654 JX440704 ID 8424 Cyrtodactylus malayanus Malaysia, Sarawak, Gunung Mulu National Park JX440655 JX440705 RMBR 00866 Cyrtodactylus malayanus Indonesia, Borneo, Kalimantan, Bukit Baka Bukit Raya GU550732 National Park ABTC 48075 Cyrtodactylus marmoratus Indonesia, Java GQ257747 TNHC 59549 Cyrtodactylus ‘‘marmoratus’’ Indonesia, Propinsi Maluku, Buru Island, Dusun Labuan JX440546 JX440604 JX440656 JX440706 n/a Cyrtodactylus mcdonaldi Australia, Northeast Queensland HQ401150 BPBM 23316 Cyrtodactylus novaeguineae Papua New Guinea, West Sepik Prov., Parkop, Toricelli Mts. JX440547 JX440605 HQ426185 HQ426274 JB 126 Cyrtodactylus oldhami captive JX440548 JX440606 JX440657 JX440707 LSUHC 8906 Cyrtodactylus pantiensis West Malaysia, Johor, Gunung Panti FR, Bunker Trail JQ889185 JX440607 JX440658 JX440708 LSUHC 8672 Cyrtodactylus paradoxus Vietnam, Hon Nghe Island JX440549 JX440608 JX440659 JX440709 CUMZ R2005.07.30.54 Cyrtodactylus peguensis Thailand, Khao Luang National Park GU550727 FMNH 236073 Cyrtodactylus philippinicus Philippines, Romblon Island JX440550 JX440609 JX440660 JQ945304 n/a Cyrtodactylus pronarus Australia, Northeast Queensland HQ401163 LSUHC 4069 Cyrtodactylus pubisulcus East Malaysia, Sarawak, Niah Cave JX4405510 JX440610 JX440661 JX440710 LSUHC 6637 Cyrtodactylus pulchellus West Malaysia, Selangor, Genting Highlands JX440711 LSUHC 6729 Cyrtodactylus pulchellus West Malaysia, Penang, Pulau Penang, Moongate Trail JX440552 JX440611 JX440662 LSUHC 4813 Cyrtodactylus quadrivirgatus West Malaysia, Pahang, Pulau Tioman, Tekek-Juara Trail JX440553 JX440612 JX440663 JX440712 KU 309330 Cyrtodactylus redimiculus Philippines, Palawan Island, Municipality of Brooke’s Point GU550740 BPBM 19731 Cyrtodactylus robustus Papua New Guinea, Sudest Island JX440554 JX440613 JX440664 JX440713 CAS 226137 Cyrtodactylus russelli Myanmar, Sagaing Division, Hkamti Township, Htamanthi JX440555 JX440614 JX440714 Wildlife Sanctuary, upper Nat E-Su stream AMS R134930 Cyrtodactylus salomonensis Solomon Islands, New Georgia I., Mt Javi, 5 km N Tatutiva JX440556 JX440615 JX440665 JX440715 Village, Marovoa 995 5 (continued on next page) 996 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003

implemented in PAUP v4.0 (Swofford, 2002). A thousand boot- ° strap replicates for each heuristic search were run with ten ran- dom sequence replicates using TBR branch swapping. The 1000 bootstrap replicates were summarized as a strict consensus tree. For ML and Bayesian analyses, the data were divided into 13 partitions, 12 corresponding to each codon position of the pro- JX440726 JX440725 JX440720 JX440721 JX440722 JX440723 JX440718 JX440719 JX440724 JX440717 RAG1 JX440716 tein-coding genes and the 13th grouping the tRNA sequences. A 5-partition scheme dividing the analyses among genes was also employed; the resulting ML and Bayesian trees had no significant differences from the 13-partition scheme, so we report only the results from the 13-partition scheme. Partitioned ML analyses were performed using RAxML HPC v7.2.3 (Stamatakis, 2006) on JX440674 JX440673 JX440669 JX440670 JX440671 JX440672 JX440668 JX440667 JX440666 PDC the concatenated dataset. Best fit evolutionary models were esti- mated in ModelTest v3.7 (Posada and Crandall, 1998) under the Akaike information criterion (Table 3). The analyses were per- formed using the more complex model (GTR + I + C) applied to all partitions due to computer programming limitations (see Ta- ble 3 for selected models). Maximum likelihood inferences were performed for 200 replicates and each inference was initiated JX440623 MXRA5 with a random starting tree. Gaps were treated as missing data and clade confidence was assessed using 1000 bootstrap pseu- doreplicates employing the rapid hill-climbing algorithm (Sta- matakis et al., 2008). Partitioned Bayesian analyses were carried out in MrBayes JX440565 JX440624 JX440564 HQ401203 JX440560JQ889189 JX440620 JX440621 JX440557 JX440617 GU366083 JX440561 JX440562 JQ889177 JX440616 JX440559 JX440619 GenBank accession numbers ND2 v3.1.2 (Ronquist and Huelsenbeck, 2003) using default priors, with models of nucleotide substitution determined in ModelTest v3.7 (Posada and Crandall, 1998)(Table 3). Two simultaneous parallel runs were performed with eight chains per run, seven hot and one cold. The analysis was run for 20,000,000 generations and sampled every 2000 generations, by which time the chains had long since reached a stationary position and the average stan- dard deviation split frequency fallen below 0.01. The program Are We There Yet? (AWTY) (Nylander et al., 2008) was employed to plot the log likelihood scores against the number of generations to assess convergence and to determine the appropriate number of burn-in trees. We conservatively discarded the first 25% of the trees as burn-in. Nodal support of 0.95 or higher was consid- ered strongly supported. In addition to analyzing the complete dataset, we also per- formed separate ML analyses of each individual locus to ensure that there were no strongly conflicting patterns among the loci. These analyses were performed in RAxML 7.2.3 under conditions similar to the combined analysis: data were partitioned by codon position (for mitochondrial data, tRNAs constituted a fourth par- Malaysia, Sabah, Poring, Sungai Kipungit trail Australia, Northeast Queensland Australia, Northeast Queensland West Malaysia, Pahang, Pulau Tioman,Papua Tekek-Juara New Guinea, Trail Misima Island JX440563 JX440622 Timor L’Est, Manufahi District, Same,Malaysia, Trilolo Johor, River Endau-Rompin, Peta, SungaiPhilippines, Kawal Palawan Island, Palawan Province,of Municipality Brooke’s Point China, Tibet Autonomous Region, Lhasa, 3Palace km WNW of Potala Thailand, Kanchanaburi Province, Sai-Yok District,Sao Ban Tha Myanmar, Sagaing Division, Alaungdaw KathapaPark, National Sunthaik Chaung (tributary to Hkaungdin Chaung) Papua New Guinea, West Sepik Prov., Parkop, Toricelli Mts. JX440558 JX440618 West Malaysia, Johor, Pulau Mentigi Locality West Malaysia, Johor, Gunung Panti FR, Bunker Trail tition), the GTR + I + C model was employed, each initial analysis was repeated for 200 replicates, and 1000 rapid bootstraps were used to assess branch support.

2.3. Divergence timing and ancestral area analyses

A timescale of evolution in Cyrtodactylus was estimated in BEAST 1.6.1 (Drummond and Rambaut, 2007). As in the preceding

sp. ‘‘Timor’’ analyses, two separate partitioning schemes were employed. However, the 13-partition analysis was terminated after 150 mil- lion generations after failing to reach convergence. Therefore, we report results of the 5-partition analysis. The analysis used a ran- Cyrtodactylus yoshii Cyrtodactylus tuberculatus Cyrtodactylus tuberculatus Cyrtodactylus tiomanensis Cyrtodactylus tripartitus Cyrtodactylus sworderi Cyrtodactylus tautbatorum Cyrtodactylus Cyrtodactylus tibetanus Cyrtodactylus tigroides Cyrtodactylus slowinskii Cyrtodactylus sermowaiensis Cyrtodactylus seribuatensis Species Cyrtodactylus semenanjungensis dom starting tree, and employed Yule tree priors and a relaxed uncorrelated lognormal clock. The analysis was run for 300 mil- lion generations, sampling every 10,000 generations, with the first ten percent of generations discarded as burn-in. Estimated

) sample sizes (>200 for all parameters) were consulted in Tracer 1.5 (Rambaut and Drummond, 2007) to ensure adequate chain length, and 95% highest posterior densities were calculated to continued ( provide credible ranges of nodal divergence dates. ZRC 2.4851 USNMFS 36138 CJS 833 LSUHC 7685 KU 309319 IRSNB 2380 LSUHC 6251 n/a USNM 579045 MVZ 233251 CAS 210205 BPBM 23317 LSUHC 6348 ID number LSUHC 8900 Three previously-used calibrations (Heinicke et al., 2011) were

Table 1 employed to date the tree. The divergence between S. roosevelti 6 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003 997

Table 2 Primers used for PCR amplification and sequencing.

Gene Primer name Primer reference Sequence

ND2 L4437b Macey and Schulte (1999) 50-AAGCAGTTGGGCCCATACC-30 L5002 Macey and Schulte (1999) 50-AACCAAACCCAACTACGAAAAAT-30 PDC PHOF1 Bauer et al. (2007) 50-AGATGAGCATGCAGGAGTATGA-30 PHOR1 Bauer et al. (2007) 50-TCCACATCCACAGCAAAAAACTCCT-30 RAG1 R13 Groth and Barrowclough (1999) 50-TCTGAATGGAAATTCAAGCTGTT-30 R18 Groth and Barrowclough (1999) 50-GATGCTGCCTCGGTCGGCCACCTTT-30 RAG1F700 Bauer et al. (2007) 50-GGAGACATGGACACAATCCATCCTAC-30 RAG1R700 Bauer et al. (2007) 50-TTTGTACTGAGATGGATCTTTTTGCA-30 MXRA5 MXRA5F2 Portik et al. (2012) 50-KGCTGAGCCTKCCTGGGTGA-30 MXRA5R2 Portik et al. (2012) 50-YCTMCGGCCYTCTGCAACATTK-30

Table 3 Best-fit models for data partitions as determined by AIC, and similar models chosen Mekong River (the greater Mekong Delta region) are excluded. for phylogenetic analyses. Eastern Indochina is defined as all of Vietnam plus those parts of Gene Model selected Model applied Cambodia and Laos east of the Mekong River. The Sunda/Wallacea ND2 region includes both Sundaland (peninsular Malaysia and islands 1st pos GTR + I + C GTR + I + C of the Malay Archipelago east to Wallace’s Line) and Wallacea (is- 2nd pos TVM + I + C GTR + I + C lands between Wallace’s Line and Lydekker’s Line). The Philippines 3rd pos GTR + C GTR + C include the entire Philippine Archipelago. The Papuan Region in- tRNAs HKY + C HKY + C cludes New Guinea, adjacent Indonesian islands east of Lydekker’s PDC Line, those parts of Melanesia inhabited by Cyrtodactylus (Bismarck 1st pos K81uf + C GTR + C Archipelago, Solomon Islands), and northeast Queensland, Austra- 2nd pos HKY + C HKY + C 3rd pos TIMef + C GTR + C lia. Several sampled taxa occur in more than one region as defined above; these were coded as occurring in C. Indochina (intermedius, RAG1 1st pos HKY + C HKY + C jarujini, oldhami, peguensis) and E. Indochina (interdigitalis), respec- 2nd pos TRN + C GTR + C tively, based on main areas of occurrence plus distributions of their 3rd pos K81uf + C GTR + C closest relatives. MXRA5 1st pos TrN + I + C GTR + I + C 2nd pos TVM + I GTR + I 3. Results 3rd pos HKY + C HKY + C The final combined dataset includes 3786 bp, of which 1808 sites are variable and 1251 are parsimony-informative. Bayesian, and S. torrei was calibrated (exponential, mean = 3, offset = 15) ML, and MP analyses of the combined dataset recover highly con- based on an amber-preserved fossil Sphaerodactylus from Hispani- cordant phylogenies (Fig. 2). Our analyses moderately support ola dated 15–20 Ma (Iturralde-Vinent and MacPhee, 1996). The (Bayesian PP/ML bootstrap/MP bootstrap = 0.74/78/37) the mono- divergence between Oedura and Woodworthia was calibrated phyly of Cyrtodactylus with one exception – the peninsular India/ (exponential, mean = 17, offset = 16) based on fossil New Zealand Sri Lanka endemic genus Geckoella is embedded within Cyrtodacty- ‘‘Hoplodactylus’’ dated to 16–19 Ma (Lee et al., 2008). The diver- lus. None of the included Palearctic bent-toed gecko genera that gence between Pygopus and Lialis was calibrated (exponential, have long been associated and confused with Cyrtodactylus, espe- mean = 10, offset = 20) based on fossil Pygopus dated to 20– cially Cyrtopodion and Mediodactylus, are part of the Cyrtodacty- 22 Ma (Hutchinson, 1998). Root height was calibrated (normal, lus + Geckoella clade (hereafter referred to Cyrtodactylus sensu mean = 200, S.D. = 13) based on estimated times of divergence of lato). However, the species C. tibetanus, which occurs north of the gekkotans from other squamates (Hugall et al., 2007; Jonniaux Himalayas and has occasionally been considered allied to some and Kumazawa, 2008; Vidal and Hedges, 2005). Palearctic bent-toed geckos (e.g. Szczerbak and Golubev, 1986; When the final timetree was obtained, ancestral biogeographic Szcerback and Golubev, 1996), is confirmed as a member of Cyrto- regions occupied by Cyrtodactylus were estimated in Mesquite 2.74 dactylus (Shi and Zhao, 2010). (Maddison and Waddison, 2011) under both MP and ML (Mk1 Basal divergences divide Cyrtodactylus sensu lato into three model) criteria. Areas were treated as unordered categorical vari- well-supported monophyletic, geographically-circumscribed ables, dividing the range of Cyrtodactylus into eight regions: Tibet, groupings: (1) C. tibetanus (Clade A in Fig. 2) of the Tibetan Plateau India/Sri Lanka, West Indochina, Thailand, Indochina, Sunda/Wall- and the northern slopes of the Himalayas; (2) a ‘‘Myanmar Clade’’ acea, Philippines, and Papua. Regions correspond to Cyrtodactylus (Clade B in Fig. 2) that includes all ten sampled Cyrtodactylus spe- faunal breaks rather than political borders, in some cases enlarged cies of diverse habits, size, and body form from Myanmar and the from traditional definitions, and are defined as follows. Tibet in- southern flank of the Himalayas, and probably additional unsam- cludes all areas to the north of the Himalayas inhabited by Cyrto- pled Cyrtodactylus from this region; and (3) a clade that includes dactylus. India/Sri Lanka includes those parts of India south of the all Geckoella plus Cyrtodactylus from Thailand, Indochina, the Malay Indo-Gangetic plain as well as the island of Sri Lanka. West Indo- Peninsula, Indonesia, the Philippines, Australia, and Melanesia, china largely corresponds to Assam and Myanmar, and is defined representing about 134 species (Clades C–M in Fig. 2). There is as the part of mainland Southeast Asia west of the Salween River, strong support from our analyses that C. tibetanus is outside all as well as the southern slopes of the Himalayas. Central Indochina remaining Cyrtodactylus + Geckoella (Bayesian PP/ML bootstrap/ largely corresponds to Thailand, and is defined as the region east of MP bootstrap = 1.00/98/78). the Salween River, north of the Isthmus of Kra, and west of the Me- Within the large third grouping, several additional geographi- kong River, except that those parts of southern Vietnam west of the cally and morphologically cohesive clades can be identified. 7 998 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003

Fig. 2. Maximum likelihood phylogeny of Cyrtodactylus, under the 13-partition scheme ( lnL 71587.112627). Nodes supported by all analyses (Bayesian PP > 0.95, ML and À MP bootstrap > 70) are indicated by stars. For nodes not supported by all analyses, support from individual analyses is indicated by black (Bayesian), gray (ML), or white (MP) circles, respectively, or support values are reported when less than 0.95 (Bayesian PP) or 70 (ML and MP bootstrap).

8 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003 999

Geckoella (Clade D) is recovered as monophyletic, but with poor relatives in Timor and New Guinea approximately 3 (5–2) Ma support (Bayesian PP/ML bootstrap/MP bootstrap = 0.9/29/–). A and 13 (18–9) Ma, respectively. significantly-supported grouping includes all sampled large- bodied species from New Guinea, adjacent island groups in Indone- sia and Melanesia, and the Cape York Peninsula of Australia 4. Discussion (‘‘Papua Clade’’; Clade I). A large, well-supported group of mainly medium-sized Cyrtodactylus (Clades J–M) includes species ranging 4.1. Phylogeny from mainland Southeast Asia to the Philippines and northwestern Australia, including one subset of species in northern Borneo and Our results support the general supposition that geographically the Philippines (‘‘Philippine Clade’’; Clade M) and another subset coherent groups of Cyrtodactylus also represent monophyletic of species from the Malay Peninsula, Greater Sundas, Sulawesi, groups. The basalmost lineage is represented in this study by only the Lesser Sundas, and the northwestern coast of Australia (‘‘Sunda C. tibetanus. It is possible that the other Tibetan/northern Himala- Clade’’; Clade J). yan species (e.g. C. medogense, C. zhaoermii; Shi and Zhao, 2010) are Separate analyses of individual loci show little significant con- also members of this clade. Members of this group have been prob- flict with one another or the phylogenies generated using the com- lematic with respect to generic allocation and have variously been bined dataset, suggesting that lineage sorting or other coalescent placed in Tenuidactylus (Szczerbak and Golubev, 1986; Szcerback processes have no effect on phylogeny estimates within Cyrto- and Golubev, 1996), Cyrtopodion (Zhao and Li, 1987), and Siwaligek- dactylus sensu lato. Of the clades described in the preceding para- ko (Khan, 2003) and some species possess characteristics that are graphs, most are recovered by either three or all four single-locus distinctive relative most other Cyrtodactylus, including small body analyses. The only exceptions are Geckoella (poor support in com- size, a clearly segmented, strongly tuberculate tail (in some spe- bined analysis; not recovered in any single-locus analysis) and cies), and large dorsal tubercles. It is possible that some unsampled Clade J (not recovered in the PDC or ND2 analyses), though all Himalayan geckos currently assigned to the genera Altigekko or Clade J species are still recovered as relatively close relatives. Addi- Indogekko also belong to this group, but each of these have puta- tionally, in comparing among the four single-locus trees, there are tively diagnostic morphological differences from Cyrtodactylus only four instances of conflict among loci, where conflict is defined (Khan, 2003). as a clade receiving ML bootstrap support >70 in one single-locus Monophyly of the Myanmar Clade is strongly supported. Sam- analysis being incompatible with a clade receiving ML bootstrap pling in this clade is especially good and even the most aberrant support >70 in another single-locus analysis. Specifically, in one member of this geographic region, C. brevidactylus—originally con- analysis C. pantiensis is recovered as most closely related to C. sidered as possibly allied to Geckoella based on morphological semenanjungensis (PDC, bootstrap support = 87), rather than C. tio- grounds (Bauer, 2002)—is embedded well within this clade. One manensis (other loci, bootstrap support = 81–90); in one analysis C. species included in our phylogeny, C. fasciolatus, belongs to this novaeguineae is recovered as most closely related to C. loriae (PDC, group as well and extends the range of the group west along the bootstrap support = 83), rather than Queensland Cyrtodactylus southern flanks of the Himalayas to northwest India. At least two (other loci, bootstrap support = 86–89); in one analysis C. ayeyarw- species, C. oldhami and C. peguensis, occur peripherally in Myanmar adyensis is outside a brevidactylus/chrysopylos/gansi clade (ND2, but are not members of this clade. The ranges of both species are bootstrap support = 88), rather than within this clade (other loci, east of the Ayeyarwady and Salween Basins and extend deep into bootstrap support = 71–75); in one analysis C. darmandvillei is Thailand; their relationships are, not surprisingly, with Thai species. recovered as most closely related to C. batucolus plus C. seribuaten- The remaining large clade of geckos includes many other geo- sis (ND2, bootstrap support = 76), rather than C. jellesmae (MXRA5, graphic groupings, some of which are exclusive. For example, all bootstrap support = 89). In each of these cases, conflicts are within Philippine Cyrtodactylus fall in a single clade (intermixed with Bor- geographically-coherent clades and do not affect biogeographic neo species) as do the sampled Geckoella, and all of the Papuan and interpretations. Queensland species (although at least one other Papuan species is The Bayesian relaxed-clock timing analysis also recovered a not a member of this clade, Oliver et al., pers. comm.). Central Indo- phylogeny that is largely concordant with the combined ML, MP, china, Eastern Indochina, and the Sunda region all harbor multiple and BI phylogenies, differing in branching pattern only at a couple lineages. One Central Indochinese clade (Clade C: C. jarujini, C. ang- poorly-supported short internal nodes. The analysis shows that ularis, C. chanhomeae) is sister to all remaining taxa, but another C. Cyrtodactylus diverged from Hemidactylus sometime near the Cre- Indochinese clade occurs elsewhere in the tree (Clade H: C. tigroides, taceous–Paleogene boundary, and that Cyrtodactylus sensu lato C. oldhami, C. peguensis), and C. interdigitalis is sister to the Sunda- has diversified throughout the Cenozoic Era (Fig. 3). Ancestral bio- land species C. elok (Clade F). Eastern Indochinese Cyrtodactylus geographic analyses depict a generally southeastward pattern of are likewise distributed in multiple clades (Clades E, F, and L, as well colonization, with Cyrtodactylus originating in the Palearctic and as C. irregularis, which is in Clade K (Fig. 2) but apparently not clo- sequentially colonizing the W. Indochina region, the C. Indochina sely related to ‘‘core’’ Clade K), each deeply divergent from their region, and the E. Indochina and Sunda/Wallacea regions; radia- respective sister taxa. Borneo supports two lineages, with three tions in Wallacea, the Philippines, and the Papua region (including species clustering with the Philippine taxa (Clade M), and three oth- Melanesia and northeastern Queensland) all trace to Sundaland ers appearing elsewhere (Clade K). Sri Lanka is represented in or ancestors. The divergence between the Palearctic species C. tibet- sample only by one species of Geckoella, with its sister taxa in pen- anus and other Cyrtodactylus occurred approximately 52 (65–40) insular India. However, Sri Lanka supports five species in the C. Ma (Fig. 3), approximately contemporaneous with the timing of fraenatus group, which on morphological grounds appears to be a the collision of the Indian and Eurasian tectonic plates (Rowley, distinct lineage, perhaps allied to the larger-bodied Myanmar taxa. 1996). The Myanmar Clade diversified starting 32 (39–23) Ma, The largest clade of Sunda/Wallacea species (Clade J) is that the divergence between the two sampled Geckoella occurred 31 sister to the main Vietnamese + Borneo/Philippine clades. This in- (38–24) Ma, and the earliest divergences in the Papuan, Sunda, cludes a subgroup endemic to the Malay Peninsula and its offshore and Philippine Clades were approximately contemporaneous, islands. This in turn is sister to C. marmoratus from Bali and Java, about 22 (28–15) Ma. The Australian species C. kimberleyensis and these are sister to a subgroup including two Malaysian species, and those in the Queensland radiation (C. tuberculatus, C. hoskini, one from Sulawesi, two from the Lesser Sundas (including Timor) C. mcdonaldi, C. adorus, C. pronarus) diverged from their closest and a newly discovered species from East Montalivet Island off 9 1000 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003

H im ala yas Asia Papuan region Phillippine region Sunda/Wallacea region E. Indochina region Pacific India/Sri Lanka region Ocean C. Indochina region W. Indochina region Tibet region

Indian Ocean

Australia

HemidactylusC. tibetanusC. slowinskiiC. russelliC. fasciolatusC. chrysopylosC. gansiC. ayeyarwadyensisC. khasiensisC. brevidactylusC. feaeC. annandaleiC. jarujiniC. angularisC. chanhomeaeC. deccanensisC. triedraC. intermediusC. hontreensisC. interdigitalisC. elokC. pulchellusC. macrotuberculatusC. tigroidesC. oldhamiC. peguensisC. loriaeC. tripartitusC. epiroticusC. louisiadensisC. salomonensisC. klugeiC. robustusC. sermowaiensisC. novaeguineaeC. adorusC. pronarusC. tuberculatusC. hoskiniC. mcdonaldiC. irregularisC. cf.C. condorensis paradoxusC. grismeriC. eisenmanaeC. marmoratusC. “marmoratus”C. quadrivirgatusC. sworderiC. semenanjungensisC. tiomanensisC. pantiensisC. darmandvilleiC. jellesmaeC. seribuatensisC. batucolusC. sp.C. “Timor” kimberleyensisC. malayanusC. cavernicolusC. consobrinusC. pubisulcusC. annulatusC. jambanganC. tautbatorumC. baluensisC. yoshiiC. aurensisC. redimiculusC. agusanensisC. phillipinnicus A B HGFEDC I K L J K M

Q 0 Ma Neogene

25 Ma

Paleogene 50 Ma

K 75 Ma

Fig. 3. Historical biogeography of Cyrtodactylus. The phylogeny is a Bayesian timetree of Cyrtodactylus, with 95% HPD intervals depicted at key nodes. Branches are colored according to the results of the ML ancestral area reconstruction. Branches where no one inferred area received at least twice as much support as the next most likely area were considered equivocal and are colored black. Other colors correspond to the map, which depicts major distributional regions of Cyrtodactylus.

the coast of the Kimberley region of Western Australia (Bauer and 2009; Shi and Zhao, 2010; Szcerback and Golubev, 1996). Not sam- Doughty, 2012). Remaining Sundaland species (Clade G) are more pling Siwaligekko, Altigekko, or Indogekko has little impact on this closely related to the Papuan Clade and some Indochinese species. biogeographic interpretation, as on morphological grounds they are certainly not embedded in core Cyrtodactylus (Clades B–M), 4.2. Biogeography and any placement for these genera still would recover a Palearctic origin for Cyrtodactylus. The subsequent spread of Cyrtodactylus to Our results depicting a Palearctic origin of Cyrtodactylus are in the east superficially resembles the pattern evident in other gekko- general agreement with biogeographic interpretations that have nid geckos with similar distributions, such as Gehyra, which origi- been made under the assumption that Himalayan bent-toed nated in Asia and colonized the southwest Pacific (Heinicke et al., geckos (including C. tibetanus and the unsampled genera Siwaligek- 2011). However, Cyrtodactylus is significantly more diverse in both ko, Altigekko, and Indogekko) represent a ‘‘transition’’ between Pale- mainland Asia and Indonesia, and species of Cyrtodactylus tend to arctic naked-toed geckos such as Cyrtopodion on one hand, and have limited ranges, whereas most Asian and Indonesian Gehyra more typical Indo-Australian Cyrtodactylus on the other (e.g., Khan, species are more widespread. 10 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003 1001

A number of fine-scale biogeographic patterns within Cyrto- these branches. One species in the Philippine/Borneo clade, C. aur- dactylus deserve comment. Our data cannot conclusively resolve ensis, occurs in the Seribuat Archipelago east of peninsular Malay- the colonization route of the peninsular Indian/Sri Lankan species sia. Two other sampled species, C. seribuatensis and C. tiomanensis, in Geckoella, as basal relationships within core Cyrtodactylus are are also endemic to this archipelago. Interestingly, none of these not fully resolved. The time of divergence post-dates the Indian/ species are closely related to one another, clearly indicating that Eurasian plate collision (Rowley, 1996), so an overland route from the South China Sea has not been a major barrier to dispersal in Myanmar was available. However, Geckoella is in a more derived Cyrtodactylus. position in the phylogeny than are the Myanmar taxa, and the In Australia, the close relationship between C. kimberleyensis ancestral area analysis does suggest it is most likely that the ances- and an undescribed Timorese species suggests that Western Aus- tor of Geckoella + Cyrtodactylus Clades E–M occupied Central Indo- tralia was colonized over water from the Lesser Sundas. Unsam- china. Thus, Geckoella may have originated via a cross-Bay of pled species from the Lesser Sundas (including C. laevigatus, C. Bengal dispersal event, which would also explain the range gap be- wetariensis and several undescribed species) are similarly small tween Geckoella and other Cyrtodactylus. (maximum 73 mm SVL) and similar in general morphology, sug- In mainland Southeast Asia, the strong geographic segregation gesting that there is a regional radiation. Not surprisingly, given of separate Cyrtodactylus clades in Myanmar, Thailand, Eastern the single species in Western Australia and its very marginal distri- Indochina, and the Malay Peninsula hints at the importance of bution, this colonization of Australia is much younger than that long-term geographic barriers in promoting regional endemism that came via New Guinea and is represented by a group of (see Fig. 1 for locations of potential barriers). The Isthmus of Kra, Queensland taxa (including C. tuberculatus), most only recently rec- separating the Thailand region from the Sunda region (which in- ognized (Shea et al., 2012). Each of these Australian colonization cludes the Malay Peninsula), has long been recognized as such a events may have been facilitated by expansion of exposed land barrier both currently as the location of an abrupt ecological tran- during periods of low sea level. Another Lesser Sunda species, C. sition from evergreen rainforest to tropical deciduous forest and darmandvillei, is related to the single species sampled from Sulaw- historically as a zone affected by marine transgressions through esi, C. jellesmae. Sulawesi supports four additional named species much of the Paleogene (Hughes et al., 2003; Woodruff, 2003; and many more remain to be described (D. Iskandar, pers. comm.). Woodruff and Turner, 2009). These transgressions could have iso- The composite geological nature of Sulawesi raises the possibility lated Thai and Sunda Cyrtodactylus radiations from one another as that it may have a compound fauna of bent-toed geckos, and sev- they diversified in the mid-Cenozoic. Members of the Myanmar eral animal clades are known to have colonized Sulawesi multiple clade occur west of the Ayeyarwady River or in isolated highland times (Stelbrink et al., 2012). Based on morphological traits, it has regions between the Ayeyarwady and Salween Rivers. These river already been suggested that the Sulawesi Cyrtodactylus fauna is valleys have likely acted as dispersal barriers due to a lack of suit- comprised of multiple distinct lineages (Iskandar et al., 2011). able habitat (most mainland Cyrtodactylus prefer karstic or other- Divergence times obtained in this study are quite similar to wise rocky terrain). Likewise, the Mekong valley may serve as a dates recovered for some Cyrtodactylus species by Siler et al. barrier between Thai and Indochinese Cyrtodactylus. An analysis (2012), even though our studies differ in calibration choice (Siler of the entire Indochinese herpetofauna suggests the Mekong is et al. calibrated the divergence between Cyrtodactylus and Gekko) not a major herpetofaunal biogeographic barrier (Bain and Hurley, and overall taxon sampling (Siler et al. included six Cyrtodactylus 2011). However, it does serve as a barrier for other groups (Meij- species). For example, we estimate that the lineage leading to C. aard and Groves, 2006) and a preference for rocky terrain separates philippinicus diverged from that leading to C. baluensis 18 (23–13) Cyrtodactylus from many other reptile and amphibian groups – the Ma, compared to an estimate of 20 (25–15) Ma in Siler et al. only Cyrtodactylus known from the Mekong valley occur in isolated (2012). Estimated mean rates of molecular evolution are also sim- rocky portions of the Mekong Delta (Bain and Hurley, 2011). Our ilar to those estimated for other gekkonids. For example, the esti- sampling of the relatively species-rich Cyrtodactylus faunas of Thai- mated mean rate of molecular evolution for RAG1 in our study 4 land, Cambodia, Laos, and Vietnam is quite limited, however, so we was 7.1 10À substitutions per site per million years, compared Ó 4 cannot discount the possibility that there is extensive geographic to 6.5 10À estimated for the genus Gehyra in a previous study Ó overlap between mainland Southeast Asian Cyrtodactylus clades (Heinicke et al., 2011). that our dataset did not capture. In the Malay Archipelago, our analyses clearly indicate that all 4.3. Taxonomy Philippine Cyrtodactylus species are closely related to species in Borneo. Unlike previous studies that have recovered a monophy- It may be argued that Cyrtodactylus is a large and still growing letic Philippine radiation (Siler et al., 2010; Welton et al., 2010a, genus that has already become unwieldy to systematists. This phy- 2010b), our tree indicates two Borneo to Philippines colonization logenetic analysis then could provide an opportunity to dismantle events in the early Neogene, about 10–20 Ma. However, support the genus in a way consistent with the well-supported monophy- values for some of the branches in the larger Philippine/Borneo letic groups hypothesized. It is clear that Geckoella is embedded group are not significant, so it is possible that a larger data set within other Cyrtodactylus and that the recognition of former as a would recover a monophyletic set of Philippine species. Within valid genus would render the latter paraphyletic. Geckoella are in- the Philippines, there is a pattern of south to north colonization, deed morphologically distinctive, with small, relatively stout in agreement with previous studies. This overall pattern is in con- bodies, short tails, and noticeably large dorsal scales, and also geo- trast to that inferred in another major Philippine gekkonid radia- graphically separated from most other Cyrtodactylus, being re- tion, Gekko. Philippine Gekko species are monophyletic, but do stricted to South Asia. However, C. brevidactylus, a member of the not have close relatives in Borneo (Rösler et al., 2011). Instead, Myanmar clade, is morphologically very similar to Geckoella. To based on molecular clock dates, Philippine Gekko most likely rafted be consistent with our tree topology recognition of Geckoella would on Palawan after rifting from the Asian mainland about 30 Ma imply the recognition of at least four new genera, one for C. tibet- (Siler et al., 2012). A similar scenario has been proposed in frogs, anus (clade A), one for the Myanmar clade (Clade B), one for some with Borneo being colonized from Palawan (Blackburn et al., Thai species (Clade C), and one for Clades J–M. Our results support 2010). While such a scenario is not compatible with the topology that C. tibetanus is more closely related to typical Cyrtodactylus and divergence dates we infer for Cyrtodactylus, we cannot wholly than to the Palearctic naked-toed clade. However, we do not know discount this possibility based on low support values for some of which, if any, other Tibetan/Himalayan geckos might belong in this 11 1002 P.L. Wood Jr. et al. / Molecular Phylogenetics and Evolution 65 (2012) 992–1003 group (Khan, 2003, 2009; Shi and Zhao, 2010; Szcerback and Golu- Lankan Cnemaspis (Reptilia: Squamata: Gekkonidae). Mitt. Mus. Naturk. Berlin bev, 1996). Further, the position of C. tibetanus as sister to all other Zool. Reihe 83, 22–32. Bauer, A.M., Doughty, P., 2012. A new bent-toed gecko (Squamata: Gekkonidae: Cyrtodactylus does not receive more than moderate support in any Cyrtodactylus), from the Kimberley region, Western Australia. Zootaxa 3187, of the analyses. The Myanmar clade receives the highest level of 32–42. support of any clade in the genus. However, morphologically, the Blackburn, D.C., Bickford, D.P., Diesmos, A.C., Iskandar, D.T., Brown, R.M., 2010. An ancient origin for the enigmatic flat-headed frogs (Bombinatoridae: Barbourula) members of this clade span almost the entire range of morpholog- from the islands of Southeast Asia. PLoS One 5, e12090. ical variation in the genus, including very large species, miniatur- Chan, K.O., Norhayati, A., 2010. A new insular species of Cyrtodactylus (Squamata: ized species, short-fingered ground dwellers, and ‘‘average-sized’’ Gekkonidae) from northeastern Peninsular Malaysia, Malaysia. Zootaxa 2389, 47–56. climbing species. Drummond, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Duran, C., Field, M., Because of these issues we advocate continued recognition of Heled, J., Kearse, M., Markowitz, S., Moir, R., Stones-Havas, S., Sturrock, S., the entire bent-toed clade as a single genus for the time being, Thierer, T., Wilson, A., 2011. Geneious v5.4. . Drummond, A.J., Rambaut, A., 2007. BEAST: Bayesian evolutionary analysis by although more detailed morphological studies that may reveal sampling trees BMC. BMC Evol. Biol. 7, 214. unambiguous synapomorphies of each of the major lineages might Grismer, L.L., Anuar, S., Quah, E., Muin, M.O., Chan, K.O., Grismer, J.L., Norhayati, A., prompt the future dismantling of Cyrtodactylus. With respect to 2010. A new spiny, prehensile-tailed species of Cyrtodactylus (Squamata: Geckoella we recommend the use of this name at the subgeneric le- Gekkonidae) from Peninsular Malaysia with a preliminary hypothesis of relationships based on morphology. Zootaxa 2625, 40–52. vel to recognize this distinctive monophyletic lineage within Cyrto- Groth, J.G., Barrowclough, G.F., 1999. Basal divergences in birds and the dactylus. Geckoella has been regularly employed as a subgeneric phylogenetic utility of the nuclear RAG-1 gene. Mol. Phylo. Evol. 12, 115–123. name in the past, so such a change is minimally disruptive (Bauer, Heinicke, M.P., Greenbaum, E., Jackman, T.R., Bauer, A.M., 2011. Phylogeny of a trans-Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single 2002, 2003; Rösler, 2000; Ulber and Gericke, 1988). There are two early colonization of Australia. Zool. Scripta 40, 584–602. other available genus-group names currently in the synonymy of Hugall, A.F., Foster, R., Lee, M.S.Y., 2007. Calibration choice, rate smoothing, and the Cyrtodactylus: Puellula Blyth 1861 (type species Puellula rubida pattern of tetrapod diversification according to the long nuclear gene rag-1. Syst. Biol. 56, 543–563. Blyth 1861) and Quantasia Wells and Wellington 1985 (type spe- Hughes, J., Round, P., Woodruff, D.S., 2003. The Indochinese–Sundaic faunal cies Hoplodactylus tuberculatus Lucas and Frost 1900). As we did transition at the Isthmus of Kra: an analysis of resident forest bird species not sample C. rubidus, a species endemic to the Andaman Islands distributions. J. Biogeogr. 30, 569–580. Hutchinson, M.N., 1998. The first fossil pygopodid (Squamata, Gekkota), and a and adjacent Cocos Islands in the Bay of Bengal, it is unclear to review of mandibular variation in living species. Mem. Queensland Mus. 41, which clade that name would apply. Quantasia could be used for 355–366. the Papuan clade (Clade I), but we do not recommend such usage Iskandar, D.T., Rachmansah, A., Umilaela, F., 2011. A new bent-toed gecko of the genus Cyrtodactylus Gray, 1827 (Reptilia, Gekkonidae) from Mount Tompotika, at this time. eastern peninsula of Sulawesi, Indonesia. Zootaxa 2838, 65–78. Iturralde-Vinent, M.A., MacPhee, R.D.E., 1996. Age and paleogeographical origin of 4.4. Conclusions Dominican amber. Science 273, 1850–1852. Johnson, C.B., Quah, E., Anuar, S., Muin, M.A., Wood, P.L., Grismer, J.L., Greer, L.F., Onn, C.K., Ahmad, N., Bauer, A.M., Grismer, L.L., 2012. Phylogeography, The phylogeny we present in this study does offer a guide to geographic variation, and taxonomy of the Bent-toed Gecko Cyrtodactylus search for synapomorphies or potential diagnostic differences quadrivirgatus Taylor, 1962 from Peninsular Malaysia with the description of a among clades. When such diagnostic traits are identified, the gen- new swamp dwelling species. Zootaxa 3406, 39–58. Jonniaux, P., Kumazawa, Y., 2008. Molecular phylogenetic and dating analyses using eric- and subgeneric-level taxonomy of Cyrtodactylus can be re- mitochondrial DNA sequences of eyelid geckos (Squamata: Eublepharidae). evaluated. In addition, it provides a scheme for outgroup selection Gene 407, 105–115. for regionally focused studies and a guide for making appropriate Khan, M.S., 2003. Questions of generic designation of angular-toed geckos of Pakistan with descriptions of three new genera (Reptilia: Gekkonidae). J. Nat. comparisons for new species descriptions. Finally, the phylogeny Hist. Wildl. 2, 1–9. we present also opens the possibility of analyzing the biogeogra- Khan, M.S., 2009. Intergeneric relations of the angular-toed geckos of circum phy, ecomorphology, and other evolutionary aspects of Cyrtodacty- Western Himalayas (Sauria: Gekkonidae). Pak. J. Zool. 41, 29–34. Kluge, A.G., 2001. Gekkotan lizard taxonomy. Hamadryad 26, 1–209. lus biology in a phylogenetic, hypothesis-driven context. For Kraus, F., 2008. Taxonomic partitioning of Cyrtodactylus louisiadensis (Lacertilia: example, long-limbed, flat-bodied cave-dwelling species occur in Gekkonidae) from Papua New Guinea. 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13 Chapter 2: A new species of lowland karst dwelling Cnemaspis Strauch, 1887 (Squamata:

Gekkonidae) from northwestern Peninsular Malaysia

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Zootaxa 3691 (5): 538–558 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3691.5.2 http://zoobank.org/urn:lsid:zoobank.org:pub:5D6AE791-06CC-447B-9DF4-E79773006EFE A new species of lowland karst dwelling Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from northwestern Peninsular Malaysia

PERRY L. WOOD, JR.1,5, EVAN S.H. QUAH2, SHAHRUL ANUAR M.S.2, 3 & MOHD ABDUL MUIN4 1Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: [email protected] 2School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia. E-mail: [email protected], [email protected] 3Center for Marine and Coastal Studies, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia 4Centre for Drug Research, Universiti Sains Malaysia, 11800 Penang, Malaysia. E-mail: [email protected] 5Corresponding author

Abstract

A new species of lowland karst dwelling Cnemaspis Strauch 1887, C. grismeri sp. nov. is described from the southeastern base of the Banjaran Bintang in northern Peninsular Malaysia. It is differentiated from its congeners by a unique combi- nation of characters including size, coloration and scalation. Cnemapis grismeri sp. nov. is most closely related to C. mcguirei, an upland species endemic to the Banjaran Bintang. This phylogeographic pattern is also seen in the upland and lowland Banjaran Bintang species of Cyrtodactylus bintangtinggi and C. bintangrendah, respectively (Grismer et al. 2012). The discovery of yet another endemic gekkonid in the poorly explored karst regions of Peninsular Malaysia under- scores the necessity for concentrated collecting efforts in these unique landscapes.

Key words: new species, taxonomy, karst, Cnemaspis grismeri, Gekkonidae, herpetofauna, Malaysia

Introduction

The gekkonid genus Cnemaspis Strauch 1887 currently contains 103 species from three non-monophyletic groups (Africa, South-Asia, and Southeast Asian [see Gamble et al. 2012; Uetz 2013]). The Southeast Asian clade (sensu Grismer et al. 2010a,b,c) comprises a group of scansorial forest dwelling lizards with numerous adaptations for moving about on flat surfaces (vegetation and rocks) during low levels of illumination. In the past, this conserved morphology and behavior lead to considerable taxonomic confusion that was ultimately disentangled with a morphological review of the entire genus and a series of new, reliable diagnostic characters (Grismer et al. 2010a,b,c). This has, in part, resulted in a rapid increase in the number of species in this genus (surpassed only by the genera Cyrtodactylus ~170 spp. and Hemidactylus ~114 spp. [Uetz 2013]) with 23 of the currently recognized 34 species having been described in the last 11 years (Chan & Grismer 2008; Chan et al. 2010; Das & Grismer 2003; Grismer 2010; Grismer & Chan 2008, 2009, 2010; Grismer & Das 2006; Grismer & Ngo 2007; Grismer et al. 2008a,b; 2009; 2010a,b,c; J. Grismer et al. 2010). Many of these new species were discovered during expeditions into previously unexplored karst forests or karst outcroppings, resulting in the acquisition of a number of new specialized lineages with highly restrictive substrate specificity, namely limestone (Grismer & Chan 2009; Grismer et al. 2008a,b, 2009, 2010a; J. Grismer et al. 2010). We report here another new lowland karst-adapted species of gecko from the limestone forests of the Lenggong Valley, Perak, in northern Peninsular Malaysia (Fig. 1). We assign the individuals of this population to the genus Cnemaspis in that they all have the unique combination of broad, flattened heads; large, somewhat forward and upwardly directed eyes with round pupils; dorsoventrally compressed bodies; and long, widely splayed limbs bearing long, inflected digits—adaptations for climbing on flat surfaces in all planes or orientations. A molecular analysis using the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) indicates they have a 7.3–11.7% sequence divergence from their closest upland relative C. mcguirei (L. Grismer et al. unpublished). Additionally, they have a unique combination of color pattern and scale characteristics that differentiate them from all other known species of Cnemaspis. Therefore, this population is described below as a new species.

538 Accepted by A. Bauer: 25 Jun. 2013; published: 24 Jul. 2013 15 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 1. Distribution map showing the location of the type locality of Cnemaspis grismeri sp. nov. in the Lenggong Valley, Perak.

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Materials and methods

Color characters were taken from digital images of living specimens cataloged in the La Sierra University Digital Photo Collection (LSUDPC) and in some cases, from living specimens. The following measurements on the type series were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate: snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90º to the inflection of the flexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90º to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eye-ball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eye-ball; eye to snout distance (ES), measured from anteriormost margin of the eye-ball to the tip of snout; eye to nostril distance (EN), measured from the anterior margin of the eye-ball to the posterior margin of the external nares; inner orbital distance (IO), measured between the anterior edges of the orbit; ear length (EL), the greatest horizontal distance of the ear opening; and internarial distance (IN), measured between the nares across the rostrum. Additional character states evaluated were numbers of supralabial and infralabial scales counted from below the middle of the orbit to the rostral and mental scales, respectively; size and number of postmental scales contacting the mental; the texture of the scales on the anterior margin of the forearm; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column (where applicable); the presence or absence of a row of enlarged, widely spaced, tubercles along the ventrolateral edge of the body between the limb insertions; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the total number of precloacal pores, their orientation, shape, and degree of separation; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales, subtibial scales, and submetatarsal scales beneath the first metatarsal; and the number of precloacal tubercles on each side of the tail base. Longitudinal rows of caudal tubercles on the non- regenerated portion of the tail are quite variable between species and useful in differentiating several taxa. Up to five pairs of the following rows may be present in varying combinations: paravertebral row — the dorsal row adjacent to the middorsal, caudal furrow; dorsolateral row — the row between the paravertebral row and the lateral, caudal furrow on the dorsolateral margin of the tail; lateral row — the row immediately below the lateral, caudal furrow; and ventrolateral row — the row below the lateral row on the ventrolateral margin of the tail. When present, this row is usually restricted to the anterior 25% (or less) of the tail. Rarely there may be a row of tubercles within the lateral, caudal furrow. Various color pattern characteristics were also evaluated (see description). Some meristic data were statistically analyzed using a one-tailed t-test with a 0.95 confidence limit on Excel 14.1.3 to test for significant differences between population means. A 1335 aligned base pair fragment of the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and its flanking t-RNAs (t-RNA-Trp, t-RNA-Ala, t-RNA-Asn, and t-RNA-Cys) was amplified using the following primers L4437b 5’-AAGCAGTTGGGCCCATACC-3’ and L5002 5’-AACCAAACCCAACTACGAAAAAT-3’ (Macey & Schulte 1999) and sequenced for nearly all species of Cnemaspis (see L. Grismer et al. unpublished). Uncorrected pairwise sequence divergences were calculated in PAUP* v4.0 (Swofford 2002) for the undescribed species (five specimens: LSUHC 9730, 9732, 9733, 9969–70) and its sister taxon (three specimens: LSUHC 8853– 8855). Specimens examined are listed in the appendix. Institutional abbreviations follow Leviton et al. (1985), except we retain ZRC (Zoological Reference Collection, Raffles Museum) for USDZ, following conventional usage. DWNP refers to the Department of Wildlife and National Parks Collection, Krau, Pahang, Peninsular Malaysia; LSUHC refers to the La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA; HC refers to the Herpetological Collection of the Universiti Kebangsaan Malaysia, Bangi, Selangor; THNHM refers to the Thailand Natural History Museum, National Science Museum, Khlong Luang, Pathum Thani, Bangkok; CUMZ refers to the Chulalongkorn University Museum of Zoology, Bangkok, Thailand; PSUZC

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refers to the Prince of Songkhla University Zoological Collection, Songkhla, Thailand; ZMKU refers to the Zoological Museum of the Kasetsart University, Bangkok, Thailand; KZM refers to the Nakhon Ratchasima Zoo Museum, ZPO, Nakhon Ratchasima, Thailand; MS refers to the collection of Montri Sumontha at the Ranong Marine Fisheries Station, Paknam, Ranong, Thailand; and UNS refers to the University of Natural Sciences, Ho Chi Minh City, Vietnam.

Systematics

Cnemaspis grismeri sp. nov. Grismer’s Rock Gecko Malaysian name: Cicak Batu Grismer Figures 2–3

Cnemaspis mcguirei Grismer, 2011:349.

Holotype. Adult male (ZRC 2.6989) collected on 7 October 2012 at Gua Asar, Bukit Kepala Gajah limestone massif, Lenggong, Perak, Malaysia (5°07.53’N, 100°58.82’E) at 78 m a.s.l. by Evan S.H. Quah and Shahrul Anuar Mohd Sah. Paratypes. ZRC 2.6990, LSUHC 9969, 9970, 9972 and 9973 were collected on 5 November 2010 from the same locality as the holotype by Evan S.H. Quah, Shahrul Anuar Mohd Sah and Mohd Abdul Muin. LSUHC 10941–10944 were collected on 7 July 2012 by the same collectors at the same locality. Diagnosis. Cnemaspis grismeri sp. nov. differs from all other Southeast Asia species of Cnemaspis in having the unique combination of adult males reaching 48.8 mm SVL, adult females reaching 50.6 mm SVL; 8–9 supralabials; 7–9 infralabials; large, lateral postmentals separated at midline by one or two smaller postmentals; forearm, subtibials, ventrals, subcaudals, and dorsal tubercles keeled; 27–32 paravertebral tubercles; tubercles on flanks, relatively small and not linearly arranged; tubercles within lateral caudal furrow; ventrolateral caudal tubercles present anteriorly; median subcaudal row not enlarged; no keeled, median subcaudal row of enlarged scales; two or three postcloacal tubercles; continuous row of eight to ten precloacal pores; subtibial scales not shield-like; no enlarged submetatarsal scales; 25–31 subdigital lamellae on fourth toe; no distinct, large, dark spots on neck; dark shoulder patch enclosing two white to yellow ocelli; prominent, wide, yellow to white, postscapular band; yellowish bars on flanks; distinct, dark, caudal bands present posteriorly; subcaudal region pigmented, not immaculate. Scalation and body size differences are summarized across all Southeast Asian species in TABLE 1. Description of holotype. Adult male; SVL 47.7 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.26), somewhat narrow (HW/SVL 0.18), flattened (HD/HL 0.44), distinct from neck; snout short (ES/HL 0.49), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum keeled, raised, larger than conical scales on occiput; prominent, supraorbital ridges; shallow frontorostral sulcus; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.21); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral slightly concave, dorsal 80% divided by longitudinal groove; rostral bordered posteriorly by supranasals and one small, azygous scale and laterally by first supralabials; eight (R,L) raised supralabials of similar size; eight (R,L) infralabials, decreasing in size posteriorly; nostrils elliptical, oriented posterodorsally; bordered posteriorly by small, granular, postnasal scales; mental large, subtriangular, bordered posteriorly by three postmentals, outer two largest; gular scales raised; throat scales larger and conical. Body slender, elongate; small, keeled, dorsal scales equal in size throughout body, intermixed with several large, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles on flanks not enlarged, moderate in size; pectoral and abdominal scales strongly keeled, raised, slightly elongate, slightly larger posteriorly; abdominal scales slightly larger than dorsals; ten continuous precloacal pores arranged in a chevron separated anteriorly by a single, non-pore bearing scale; precloacal depression present; forelimbs moderately long, slender; dorsal scales of brachium raised, keeled; dorsal scales of forearm same as brachials; ventral scales of brachium weakly keeled, raised, juxtaposed; scales beneath forearm, weakly keeled, raised; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath

A NEW SPECIES OF CNEMASPIS FROM MALAYSIA Zootaxa 3691 (5) © 2013 Magnolia Press · 541 18 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing weak; fingers increase in length from first to fourth with fourth and fifth equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing present; toes increase in length from first to fourth with fourth and fifth equal in length; 29 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; caudal scales raised, keeled, juxtaposed anteriorly; shallow middorsal furrow; deeper, single lateral furrow; no enlarged, median subcaudal scales; subcaudals keeled; no median row of enlarged keeled subcaudal scales; transverse tubercle rows do not encircle tail; caudal tubercles present in lateral furrow; three enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail; tail 1.45% of SVL.

FIGURE 2. Upper left: adult female Cnemaspis grismeri sp. nov. paratype LSUHC 9973 in dark daytime coloration. Upper right: adult male C. mcguirei LSUDPC 5175. Lower left: adult male C. grismeri (uncataloged) sleeping on a leaf in its light, nighttime color phase. Lower right: adult male C. grismeri sp. nov. LSUHC 9972 in dark daytime coloration.

Coloration (in life). Dorsal ground color grey to brown; head and body overlain with irregularly shaped, small, dark and yellowish flecks giving an overall mossy appearance; cream to yellowish markings on top of head; thin dark postorbital stripe extending onto nape; paired, elongate, medial, yellowish markings on nape followed by small, indistinct, black shoulder patches enclosing two yellow ocelli; ocellus dorsal to forelimb insertion distinct and another anterior to forelimb insertion very weak; shoulder patch edged posteriorly by wide, postscapular band that is yellow laterally and white medially; irregularly shaped, offset, paravertebral, yellowish markings on dorsum extend to base of tail; distinct, transversely elongate, yellow bars on flanks; diffuse brown and dull yellow bands encircle tail anteriorly; posterior portion of tail with dark brown and off-white bands ; irregularly shaped yellowish to dull white markings on limbs; dark and light diffuse bands encircling digits; ventral surfaces of head, body, and limbs dull beige, immaculate, darkening laterally; subcaudal region suffused with pigment, not immaculate.

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Variation (Fig. 2,3). The overall general color pattern of the paratypes closely approaches that of the holotype. The postscapular band in the paratype LSUHC 9972 is offset. The light dorsal markings in ZRC 2.6990, LSUHC 9969 and 9973 appear more as transverse bands than paravertebral blotches. The black shoulder patch and postscapular bar in females is not nearly as well developed as it is in males (Fig. 3). LSUHC 10943 is missing a tail and the tails of LSUHC 9969–70 are mostly or entirely regenerated. Meristic differences are presented in TABLE 2. During the evening, the color pattern lightens considerably (Fig. 2).

FIGURE 3. Differences between male and female Cnemaspis grismeri sp. nov. in the development of the dark shoulder patch. Upper: uncataloged adult female. Lower: adult male holotype ZRC 2.6989.

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Commercial sale or deposition in a public library or website is prohibited. chanthaburiensis chanthaburiensis 1 1 0

23–25 chanardi chanardi ed on the next page

1

6–8 7–9 40.1 42.1 caudanivea caudanivea 00 00 0 ……continu

47.1 weak 1 1

ghlighted values are discretely boulengeri boulengeri /0 0 /0 1 0 0 0–3 0 0

32–38 20–27 20–30 biocellata biocellata // 1 40.1 69 21–23

le. Ant=anteriorly only. Hi bayuensis bayuensis

8,9 5–7 6,7 7,8 4–8 7–9 46.1 9,10 6–10 8,9 8,9 7–10 8–10 baueri baueri 00 0 11 1 0 5–9 8–12 64.9 11–13 10–12

aurantiacopes aurantiacopes / / 0 0 / 0 0 0 1 000 0 0

56.6 9–11 8–10 argus argus . Cnemaspis /=Character not applicab 8 8 1 8–10 28–32 23–29 18–27 23–30

affinis affinis 0 0 1 1 1 1 1 1 00 00 0 48 65.2 5,6 9–13 8–10 20–28 sp. nov. C. grismeri C. Southeast Asian species of from those values in those values from s continuous (1) or separated (0) s elongate (1) or round (0) 0 0,1 / / 0 sent (1) or absent (0) on flanks Diagnostic Diagnostic characters of s Maximum SVL Supralabial TABLE 1. diagnostic and differ differ diagnostic and Tubercles of ventralmost row on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely spaced and randomly more distributed (0) Caudal tubercles in lateral furrow (0) (1) or not No. of paravertebral tubercles Precloacal pore Precloacal pore Tubercles pre Infralabials Ventral scales keeled (1) or (0) not No. of precloacal pores

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chanthaburiensis chanthaburiensis

0 0 0 0 0 0 0

1–3 chanardi chanardi 0 1 1 1 0 1 0 0 weak 25–30 25–29

caudanivea caudanivea

1 0 1 0 1 0 0 0 0 0 0 0 0 ……continued on the next page 1,2 23–30

boulengeri boulengeri 0 1 1 1 1 1 0 1 1 0 1 29–32

biocellata biocellata

0 0 0 1 1 0 bayuensis bayuensis

1 0 0 0 2 baueri baueri

0 1 0 0 0 0 0 0 0 0 0 0 0 0 aurantiacopes aurantiacopes 1 1 0 0 0 0 0 1 1 1 27–30 26–27 27–30 29–37

argus argus 0 0 32–35

affinis affinis

1 1 2 1–4 2 2 0 0 0 1 1 1 1 1 0 0 0 0 0 0 0 17–20 (Continued) (Continued)

Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 1 1 Ventrolateral caudal tubercles anteriorly(1) not or (0) TABLE 1. Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (1) (0) or not Single rowmedian of keeled subcaudals (1) not or (0)Caudal tubercles encircle tail (1) or (0) not 0 Enlarged subcaudalmedian scale row (1) or not (0) No. of postcloacal tubercles 0 scales Enlarged present femoral (1) or absent (0) Shield-like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) (0) or not 0 0 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 0 0 0 0 No. of 4th toe lamellae

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limi

/ 0 1 0

7–9 88.2 8–11

laoensis ed on the next page /

9 1 7 0

40.9

kumpoli

1 1 1 0 / / 0 0 63 ……continu 7,8 7,8 / 6–8 29–35 22 25–33

kendallii / 0 0 0 58

weak 10–12

karsticola 0 0 0 0 0 1 0 / 0 1 1 7,8 48.1 17–19 20–26

kamolnorranathi 1 1 1 1 0 8,9 7,8 6,7 37.8 19–24

harimau 1 0 9,10

flavolineata

1 1 0 1 1 0-weak

8–10

flavigaster 1 0 0 0 0 0 1 1

7,8 7 4 9,10

21–24 22–24 18–20

dringi

1 0 0 0 0 0 0 0 0 9 8–10 8–10 9,10 7,8 6,7 7–9 11 5,6 46.4 50.1 46.7 40.7 25–27

(Continued) Tubercles of ventralmost row on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely spaced and randomly more distributed (0) Tubercles present (1) or absent (0) on flanks Precloacal pores elongate (1) or round (0) No. of paravertebral tubercles Caudal tubercles in lateral furrow (0) (1) or not Infralabials Ventral scales keeled (1) or (0) not No. of precloacal pores Precloacal pores continuous (1) or separated (0) 0

Supralabials Maximum Maximum SVL TABLE 1.

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limi 1 0 0 0

weak 30–35

laoensis ed on the next page 0 0 0 1 29

kumpoli 0 0 0 0 0 0 0 0 0 0 0 / 0 0 1 0 ……continu 2,3 2,3 1,2 34–41

kendallii 1 0 1 0 1 1 1 0

karsticola 1 0 0 0 0 0 1 1 1 1 1

kamolnorranathi 0 0 0 0 1 0 0

weak

harimau 0 0 1 0

flavolineata 0 1 1 weak 1 1 1 1 1 0 0 / 1 0 0 0 0 0 0 0 0

flavigaster 0 1 0 0 0 0 1 0 0 0 0 0

dringi 0 1 2 1,2 2,3 0 1,2 2,3 1,3 0 0 1 1 1 1 1 0-weak 32–35 29–34 23–26 25–30 24–28 27–30 23–36

(Continued)

Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (0) (1) or not Single rowmedian of keeled subcaudals (1) not or (0) 1 0 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 Ventrolateral caudal tubercles anteriorly (1) not or (0) 0 TABLE 1. Caudal tubercles encircle tail (1) or (0) not Enlarged subcaudalmedian scale row (1) or not (0) Enlarged femoral scales Enlarged present femoral (1) or absent (0) 0 0 0 No. of postcloacal tubercles Shield-like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) (0) or not 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 No. of 4th toe lamellae

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pseudomcguirei 9 1 0 1 0 ant 1–5 42.5

perhentianensis 0 0 1 0 0 0 1 1 47 7–8 8,9 6–8 8–10 22–27 26–32

pemanggilensis / 0 1 0 0 1 76

……continued on the next page 10–13 10–14

paripari 0 1 1 0,1 1 0 0 0 12 2–6 10,11 26–31 30–37

nuicamensis 0 1 0 0 0 1 1 7–9 4–6 47.5 50.7

niyomwanae 0 0 3 0 0 0 56.8 8–11

nigridia 0 1 0 0

10,11 12–14 39–43 26–31 20–22

neangthyi / / 0 1,0 0 0 / / / / 1 1 1 0 1 1 1 1 1 1 0 2 54 75.5

monachorum 0 0 0 3 1 0 7,8 11–13 5–7 11–12 9–11 6–8 6–7 32.9 11–19

mcguirei

1 0 0 1 1 0 0 0 1 0 65 7,8 7–9 5–10 26–32 (0) (0) in lateral furrow (1) or not or (1) furrow lateral in (Continued) (Continued) Maximum SVL Supralabials TABLE 1.

Infralabials Ventrolateral caudal tubercles anteriorly(1) not or (0) Lateral caudal tubercle row present (1) or absent (0) 1 ant Ventral scales keeled (1) or (0) not No. of precloacal pores Precloacal pores continuous (1) or separated (0) 0 Precloacal pores elongate (1) or round (0) No. of paravertebral tubercles Tubercles of ventralmost row on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely space and more randomly distributed (0) Tubercles present (1) or absent (0) on flanks Caudal tubercles

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pseudomcguirei 0 1 2,3 0,1

perhentianensis 0 0 0 0 0 1 28–31 23–25

pemanggilensis 1 1 2 3–4 0 0 0 ……continued on the next page

paripari 1 0 1 1 1 0 0 0 0 1 1 26–31 27–31

nuicamensis 0 0 0 1 1 1 0

weak weak

niyomwanae 0 0 1 0 0 0

nigridia 0 1 1 1

neangthyi / 0 0 / 2–4 1,2 2–4 2 1 0 0 0

monachorum 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0

0 1 1 0 mcguirei mcguirei 2,3 1,2 30–35 25–27 22–25 28,29 31–34 29–33 (Continued) (Continued) Single rowmedian of keeled subcaudals (1) not or (0) 0 0 0 0 0 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (0) (1) or not TABLE 1. Caudal tubercles encircle tail (0) (1) or not No. of postcloacal tubercles

Enlarged subcaudalmedian scale row (1) or not (0) 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 Shield-like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) (0) or not 0 0 1 0 0 0 0 0 0 0 Enlarged femoral scales Enlarged presentfemoral (1) or absent (0) 0 0 No. of 4th toe lamellae

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grismeri

1 1 8,9 8–10 27–32

punctatonuchalis / 0 1 1 8 0 0 49.6 50.6 24–27

huaseesom

0 0 0 0 0 1 0 ……continued on the next page

5,6 6–8 7,8 7–9 7–9 37.9

18–24

narathiwatensis 0 0 0 1 1 1 1 1 1 ant 1 1 7–9 3–6 43.2 9,10 28,29

vandeventeri 1 0 0 0 0 1 4 1 8,9 25–29

tucdupensis

0 1 0 1 0 0 51 44.7 8–10 18–21

siamensis

1 0 0 0 0 8,9 6–8 7,8 7–9 39.7 19–25

shahruli / / / 0 0 / / / 0 0 0 / 0 / 1 0 8–10 10,11 19–23

roticanai

1 1 0 1 0 1 0 1 1 1 1 47 36.5

7,8 8,9 3–6

25–27

psychedelica

/ 0 0 0 0 0 0 0 0 0 0 7,8 5–7 75.3 34–48 (1) or not (0) udal tubercles anteriorly (Continued) (Continued)

No. of paravertebral tubercles Tubercles present (1) or absent (0) on flanks Caudal tubercles in lateral furrow (0) (1) or not Ventrolateral ca Infralabials Maximum SVL Supralabials TABLE 1. Lateral caudal tubercle row present (1) or absent (0) Precloacal pores elongate (1) or round (0) Tubercles of ventralmost row on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely space and more randomly distributed (0) Ventral scales keeled (1) or (0) not No. of precloacal pores Precloacal pores continuous (1) or separated (0) / 0

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grismeri grismeri

0 1 0 1

punctatonuchalis 1 0 1 0 0 0 0 0 1,3 2,3

29 25–31

huaseesom 0 0 0 0 0 0 0 1,2

21–31

narathiwatensis

1 0 0 0 0 0 0 1 2,3 24–26

vandeventeri

0 0 1 0 0 0 0 0 1 1–3 weak 24–28

tucdupensis

1 0 0 0 0 0 0 1 0 0 0 0 0 27–30

siamensis

0

weak 1

shahruli 0 0 1 1 1 0 0 1 1 1–3 1,2 0–3 21–30 24–26

roticanai 0 1 1 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 1,2 26–29

psychedelica

0 1 0 1 1 weak 1 0 1 1,2 25–28

(Continued) (Continued)

Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (0) (1) or not TABLE 1. Single rowmedian of keeled subcaudals (1) not or (0) Enlarged subcaudalmedian scale row (1) or not (0) No. of postcloacal tubercles Caudal tubercles encircle tail (0) (1) or not Enlarged femoral scales Enlarged present femoral (1) or absent (0) 1 Shield-like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) (0) or not Enlarged submetatarsal scales on 1st toe (1) or not (0) No. of 4th toe lamellae

A NEW SPECIES OF CNEMASPIS FROM MALAYSIA Zootaxa 3691 (5) © 2013 Magnolia Press · 551 28 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. 10944 LSUHC paratype s for character 10943 LSUHC paratype ……continued on the next page 10942 LSUHC paratype 10941 LSUHC paratype 9973 LSUHC paratype 9972 LSUHC paratype ZRC ZRC 2.6990 paratype il; r=regenerated tail; /=unable to assess. See Materials and method 9970 LSUHC paratype 9969 LSUHC paratype 1 0 1 0 / 1 1 / 0 / 1 1 / 0 / 1 1 1 / 1 0 0 1 1 1 1 0 1 0 1 / 1 / 1 / / 0 1 0 / 1 1 0 1 0 1 / 1 1 1 / 1 0 1 1 1 / 1 1 / 0 1 1 / 1 1 / 0 1 1 1 1 0 1 / / / 1 1 1 8 8 9 9 9 8 8 8 9 9 8 7 8 7 8 7 8 8 9 8 M F F F M F M F F F 10 / / / 8 / 9 / / / 27 31 27 30 32 27 28 30 30 31 ZRC ZRC 2.6989 holotype

abbreviations. No. of precloacal pores Precloacal pores in a continuous row (1) or separated (0) medially Precloacal pores elongate (1) or round (0) No. paravertebral tubercles Tubercles of ventral row most on flank linearly arranged and in contact or nearly so (1) or tubercles on flank widely space and more randomly distributed (0) Caudal tubercles in lateral caudal furrow (1) (0) or not Ventrolateral caudal tubercles anteriorly (1) or not (0) Lateral caudal tubercles present (1) or not (0) Sex Supralabials Infralabials Ventral scales keeled (1) or smooth (0) Tubercles present absent (1) (0) or on flanks TABLE 2. Morphological data taken for the type series. BT=broken ta Cnemaspis grismeri

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10944 LSUHC paratype 10943 LSUHC paratype ……continued on the next page 10942 LSUHC paratype 10941 LSUHC paratype 9973 LSUHC paratype 9972 LSUHC paratype ZRC ZRC 2.6990 paratype 9970 LSUHC paratype 9969 LSUHC paratype 0 1 0 0 0 1 0 0 / 3 0 / 0 0 / 0 0 2 / 1 1 0 / 0 0 0 0 2 0 1 1 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 0 0 0 0 0 3 0 1 0 1 1 0 0 0 0 0 0 0 0 0 3 1 0 0 1 1 0 0 0 0 0 0 / 0 0 3 1 0 0 / 1 0 0 0 0 0 / 0 0 0 2 0 1 / 1 1 0 0 0 0 / 0 0 0 3 1 0 0 1 0 0 0 0 0 0 0 2 1 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 0 1 0 0 0 1 0 29 30 29 29 30 28 25 30 27 31 ZRC ZRC 2.6989 holotype 1 0 0 0 0 0 0 0 0 0

(Continued) (Continued) Dark, longitudinal, gular markings present (1) or absent (0) Single median row of keeled subcaudals or (1) not (0) Caudal tubercles encircle tail (1) or not (0) Enlarged median subcaudal scale row not (1) (0) or No. of postcloacal tubercles scales presentEnlarged femoral (1) or absent (0) Shield-like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) (0) or not Enlarged submetatarsal scales on first toe (1) (0) or not No. of 4th toe lamellae Head and tail yellow in (1)males or not (0) to anterior crescent yellow to White not (1) (0) or forelimb Black shoulder patch enclosing (0) not or (1) ocelli white to yellow Distinct, large, isolated, dark spots on neck (1) (0) or not Distinct, dark and light caudal bands present (1) or absent (0) Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (1) (0) or not TABLE 2.

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d i v a r 10944 LSUHC paratype g d i v a r 10943 LSUHC paratype 10942 LSUHC paratype 10941 LSUHC paratype 9973 LSUHC paratype 9972 LSUHC paratype ZRC 2.6990 paratype 9970 LSUHC paratype 9969 LSUHC paratype g 4.9 7.7 5.4 8.2 4.8 8.7 8.5 4.5 5.4 7.1 5.2 8.2 2.6 8.4 5.9 3.4 4.8 8.3 2.3 6.1 7.7 5.4 3.7 5.2 4.3 7.3 2.8 6.6 7.7 3.8 4.8 3.7 5.4 8.5 5 1.1 2.7 6.4 8.1 5.3 3.1 1.2 3.5 4.8 7.7 2.7 5.1 1.3 6 7.2 5.1 2.6 3.6 1.3 8.1 5.2 2.5 4.5 1.2 5.7 6.6 2.8 8.1 3.4 1.3 8.1 2.6 4.9 1.2 5.1 6.1 3.5 3.4 1.1 8.1 2.5 4.6 1.3 5.1 6.1 2.5 3.6 1.2 2.5 8.1 5.8 1.2 6.1 2.7 3.6 5.3 1.2 4.5 1.1 2.5 6.1 2.6 1.2 3.5 4.5 1.1 2.9 1.3 6.1 1.1 4.9 1.3 2.9 1.2 1.5 47.7 69.3 50.6 54.2r 48.6 49r 10.1 43.5 20.1 21.2b 10.6 48.8 12.4 21.2 47.9 9.8 45.5 13.1 20.2 64 45.7 9.7 12.7 19 61.1 48.5 10.6 12.1 24.3 70.1 44.4 9.7 13 20.1 BT 10.1 48.2 12.4 20.3 68.1 10.1 12.6 22.7 9.4 12.6 20.1 11.8 10.1 22.1 12.6 ZRC 2.6989 holotype

(Continued) IO IO EL IN ED EE ES EN AG AG HL HW HD TL TW FL TBL SVL SVL TABLE 2.

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Distribution. Cnemaspis grismeri sp. nov. is known from Gua Asar at the Bukit Kepala Gajah limestone massif in Lenggong Valley, Perak (Fig. 1). This species has also been observed at other caves, namely Gua Kijang, Gua Puteri and Gua Ngaum that are also part of the Bukit Kepala Gajah limestone massif as well as its surrounding karst walls and karst forest. Natural history. Cnemaspis grismeri sp. nov. is a lowland species found on the inner walls of limestone caves such as Gua Asar and Gua Kijang and near cave entrances and on the karst walls outside of caves as well as outcroppings in the lowland karst forest surrounding limestone massifs (Fig. 4). Cnemaspis grismeri sp. nov. is diurnal but is only observed crawling about in cracks and in the shadows of the karst boulders especially in vertical crevices. The lizards are very wary and quickly retreat deeper into their refuges when approached. Their behavior is similar to that of its upland closest relative C. mcguirei that behaves the same way on granite boulders at Bukit Larut, Perak (Grismer 2011). At night the color of C. grismeri sp. nov. lightens and lizards have been observed sleeping on the leaves of low vegetation near the karst walls and on vines and the aerial roots of fig trees (Ficus sp.) next to the limestone walls (Fig. 2 & 4). Specimens LSUHC 10942–43 are gravid females, which indicate that breeding takes place around the month of July.

FIGURE 4. Microhabitat structure of the karst formations for Cnemaspis grismeri sp. nov. from Lenggong Valley, Perak.

Etymology. This species is named in honor of Prof. Dr. L. Lee Grismer of La Sierra University, Riverside, California, USA for his tremendous contributions to the advancement of the field of herpetology in Malaysia. Coincidentally, Cnemaspis grismeri sp. nov. is the closest relative of C. mcguirei, a species that Dr. Grismer had named in honor of his close friend and colleague Prof. Dr. Jimmy A. McGuire of the University of California Berkeley, USA. Comparisons. Cnemaspis grismeri sp. nov. differs from all other species of Cnemaspis in numerous aspects of scalation and color pattern as summarized in TABLE 1. It is most similar to its sister species C. mcguirei (L. Grismer et al. unpublished) but differs in being smaller (maximum SVL=50.6 mm in C. grismeri sp. nov. versus

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SVL=65.1 mm in C. mcguirei) and having a significantly lower mean number of subdigital lamellae on the fourth toe (28.8 versus 32.5; p<0.01). Additionally, the lower, anterior shoulder spot is less prominent and the lateral baring is generally more distinct (Figs. 2 & 3). Cnemapsis grismeri sp. nov. is the sister species of C. mcguirei who are separated by a 7.3–11.7% sequence divergence in the mitochondrial gene ND2.

Discussion

Grismer (2011) included Cnemaspis grismeri sp. nov. as part of C. mcguirei but noted that the trenchant differences in body size and substrate preference (karst for C. grismeri sp. nov. and granite for C. mcguirei) might indicate that these are related but distinct lineages. Preliminary genetic work on the entire genus Cnemaspis (Grismer et al. unpublished) utilizing the mitochondrial ND2 gene demonstrated that they are sister species, but have a sequence divergence from one another of 7.3–11.7% which is also observed between other sister species groups. Additional differences in color pattern and scalation support the hypothesis that these populations are separate species (TABLE 1). A phylogeographic pattern of upland and lowland sister species within the Banjaran Bintang is beginning to emerge. Grismer et al. (2012) reported the same pattern for the upland Cyrtodactylus bintangtinggi (Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wnagkulangkul, Grismer, Pauwels) and the lowland Cyrtodactylus bintangrendah (Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wnagkulangkul, Grismer, Pauwels)—species that are syntopic with C. mcguirei and C. grismeri sp. nov., respectively. The karst regions of Peninsular Malaysia are proving to be warehouses of species diversity and endemism, especially with respect to gekkonids (Grismer 2011; Grismer & Chan 2009; Grismer et al. 2008a,b; 2009, 2010a,c, 2012; J. Grismer et al. 2010). The discovery of yet another endemic gekkonid in these poorly explored areas underscores the need for concentrated collecting efforts in these unique landscapes. Based on the latest assessment of the distribution of karst in Peninsular Malaysia (Price 2005) we estimate we have only explored 5% of what is available. Additional field work in these unexplored karst formations is needed to help better understand the herpetofaunal diversity in Peninsular Malaysia.

Acknowledgements

For assistance in the field we are indebted to Mr. Yusof Omar. We are also grateful to Prof. Dr. L. Lee Grismer of La Sierra University, Riverside, California, USA, for his valuable help and mentoring. This research was supported in part by a Universiti Sains Malaysia grant 815075 to Shahrul Anuar. EQSH’s research is supported in part by the USM Fellowship Scheme.

References

Chan, K.O. & Grismer, L.L. (2008) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Selangor, Peninsular Malaysia. Zootaxa, 1877, 49–57. Chan, K.O., Grismer, L.L., Shahrul, A., Quah, E., Muin, M.A., Savage, A.E., Grismer, J.L., Norhayati, A., Remegio, A.C. & Greer, L.F. (2010) A new endemic rock Gecko Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia. Zootaxa, 2576, 59–68. Das, I. & Grismer, L.L. (2003) Two new species of Cnemaspis Strauch 1887 (Squamata, Gekkonidae) from the Seribuat Archipelago, Pahang and Johor States, West Malaysia. Herpetologica, 59, 544–552. http://dx.doi.org/10.1655/02-22 Dring, J.C.M. (1979) Amphibians and reptiles from northern Trengganau, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum (Natural History), 34, 181–241. Gamble, T., Greenbaum, E., Jackman, T.R., Russell, A.P. & Bauer, A.M. (2012) Repeated origin and loss of adhesive toepads in geckos. PLoS ONE, 7, e39429. http://dx.doi.org/10.1371/journal.pone.0039429 Grismer, J.L., Grismer, L.L. & Chav, T. (2010) New Species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Southwestern Cambodia. Journal of Herpetology, 44, 28–36. http://dx.doi.org/10.1670/08-211.1

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Grismer, L.L. (2010) The first record of the genus Cnemaspis Strauch (Squamata: Gekkonidae) from Laos with the description of a new species. Zootaxa, 2475, 55–63. Grismer, L.L. (2011) Lizards of Peninsular Malaysia, Singapore and their Adjacent Archipelagos. Edition Chimaira, Frankfurt, 728 pp. Grismer, L.L & Chan, K.O. (2008) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Zootaxa, 1771, 1–15. Grismer, L.L. & Chan, K.O. (2009) A new species of karst dwelling Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Sarawak, Borneo. Zootaxa, 2246, 21–31. Grismer, L.L. & Chan, K.O. (2010) Another new rock gecko (genus Cnemaspis Strauch 1887) from Pulau Langkawi, Kedah, Peninsular Malaysia. Zootaxa, 2419, 51–62. Grismer, L.L. & Das, I. (2006) A new species of gekkonid lizard of the genus Cnemaspis Strauch 1887 from Pulau Pemanggil, Johor, West Malaysia. Herpetological Natural History, 10, 1–7. Grismer, L.L. & Ngo, V.T. (2007) Four new species of the gekkonid genus Cnemaspis Strauch 1887 (Reptilia: Squamata) from Southern Vietnam. Herpetologica, 63, 482–500. http://dx.doi.org/10.1655/0018-0831(2007)63[482:FNSOTG]2.0.CO;2 Grismer, L.L., Chan K.O., Nurolhuda, N. & Sumontha, M. (2008a) A new species of karst dwelling gecko (genus Cnemaspis Strauch 1887) from the border region of Thailand and Peninsular Malaysia. Zootaxa, 1875, 51–68. Grismer, L.L., Chan, K.O., Quah, E., Muin, M.A., Savage, A.E., Grismer, J.L., Norhayati, A., Greer, L.F.III, & Remegio, A.C. (2010c) Another new, diminutive Rock Gecko (Cnemaspis Strauch) from Peninsular Malaysia and a discussion of resource partitioning in sympatric species pairs. Zootaxa, 2569, 55–66. Grismer, L.L., Grismer, J.L., Wood, Jr., P.L. & Chan, K.O. (2008b) The distribution, taxonomy, and redescription of the geckos Cnemaspis affinis (Stoliczka 1887) and C. flavolineata (Nicholls 1949) with descriptions of a new montane species and two new lowland, karst- dwelling species from Peninsular Malaysia. Zootaxa, 1931, 1–24. Grismer, L.L., Ngo, V.T. & Grismer, J.L. (2010b) A colorful new species of insular rock gecko (Cnemaspis Strauch 1887) from southern Vietnam. Zootaxa, 2352, 46–58. Grismer, L.L., Norhayati, A., Chan, K.O., Belabut, D., Muin, M.A., Wood, Jr., P.L. & Grismer, J.L. (2009) Two new diminutive species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Peninsular Malaysia. Zootaxa, 2019, 40–56. Grismer, L.L., Sumontha, M., Cota, M., Grismer, J.L., Wood, Jr., P.L., Pauwels, O.S.G. & Kunya, K. (2010a) A revision and redescription of the rock gecko Cnemaspis siamensis (Taylor 1925) (Squamata: Gekkonidae) from Peninsular Thailand with descriptions of seven new species. Zootaxa, 2576, 1–55. Grismer, L.L., Wood, Jr., P.L., Quah, E.S.H., Shahrul, A., M.A. Muin, Sumontha, M., Norhayati, A., Bauer, A.M., Wangkulangkul, S., Grismer, J.L. & O.S.G. Pauwels (2012) A phylogeny and taxonomy of the Thai-Malay Peninsula Bent-toed Geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species. Zootaxa, 3520, 1–55. Leviton, A.E., Anderson, S.C., Gibbs, R.H., Heal, E. & Dawson, C.E. (1985) Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, 802–832. Macey, J. & Schulte, J. (1999) Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution, 12, 250–272. http://dx.doi.org/10.1006/mpev.1999.0615 Price, L. (2005) Geology of Peninsular Malaysia. In: Geyer, E., Schmidt, F. & Jeutter, P. (Eds.), Expedition Gunung Lanno, Malaysia. Verein für Höhlenkunde in Obersteier, Bad Mitterndorf, Austria, pp. 30–31. Swofford, D.L. (2002) Paup*: Phylogenetic Analysis Using Parsimony (and Other Methods), Version 4.0. Sinauer Associates, Sunderland, Massachusetts. Uetz, P. & Hošek, J. (eds.) (2013) The Reptile Database, Available from: http://www.reptile-database.org (Accessed 23 July 2013)

APPENDIX. The following specimens were examined.

Cnemaspis affinis (Stoliczka): LSUHC 6695, 6758–59, 6773, 6788, 8975, Pulau Penang, Penang, Malaysia. Cnemaspis aurantiacopes Grismer & Ngo: LSUHC 9528–29, 9531, 9535, 9539, Kien Giang Prov., Hon Dat Dist., Hon Dat Hill, Vietnam. Cnemaspis baueri Das & Grismer: ZRC 2.5291 (holotype), ZRC 2.5292–99, LSUHC 3921–24, 4700–01, 4717–29, 4744, 4808, 7272–74, 7301–03, 7319, Pulau Aur, Johor, Malaysia. Cnemaspis bayuensis Grismer, Grismer, Wood & Chan: LSUHC 9070–74, Gua Bayu, Kelantan, Malaysia. Cnemaspis biocellata Grismer, Chan, Nurolhuda & Sumontha: ZRC 2.6693–98, Kulau Perlis, Perlis Malaysia; MS 30, Khao Tohphayawang, Muang Satun, Satun Province, Thailand. Cnemaspis boulengeri Strauch: CAS 73745, LSUHC 9542, 9578–79, MCZ 39014-23, Pulo Condore (= Con Dao), Vietnam. Cnemaspis caudanivea Grismer & Ngo: LSUHC 9544–48, Kien Giang Province, Kien Hai District, Hon Tre Island, Vietnam. Cnemaspis chanthaburiensis Bauer & Das: FMNH 215979 (holotype) and FMNH 191479 (paratype), Khao Soi Daouw (Dao) Wildlife Sanctuary, Pongnomron (Pong Nam Ron) District, Chanthaburi Province, Thailand; BMNH 1917.5.14.4

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(paratype), Chantaburi Province, Thailand; FMNH 215978 (paratype), Khao Khiew (Khieo) Wildlife Sanctuary, Chon Buri Province, Thailand; FMNH 215980 (paratype), Suan Kaset, Muang District, Chantaburi Province, Thailand; LSUHC 7882, Phnom Samkos, Pursat Province, Cardamom Mountains, Cambodia. Cnemaspis dringi Das & Bauer: FMNH 148588 (holotype), Labang Camp (03º 20’ N; 113º 29’ E), Bintulu District, Fourth Division, Sarawak, Malaysia; FMNH 221478 (paratype), Sungai Segaham, Belaga District, Seventh Division, Sarawak, Malaysia. Cnemaspis flavigaster Chan & Grismer: HC 0082–86, ZRC 2.6708–11, LSUHC 8835–36, Forest Research Institute Malaysia, Kepong, Selangor, Malaysia; BM 1898.9.22.216, Batu Caves, Selangor, Malaysia. Cnemaspis flavolineata (Nicholls): LSUHC 8097, Frazer’s Hill, Pahang, West Malaysia; LSUHC 9110, 9159–60, HC 303, Cameron Highlands, Pahang, West Malaysia. Cnemaspis karsticola Grismer, Grismer, Wood, & Chan: LSUHC 9053–56, Gunung Reng, Kelantan, Malaysia. Cnemaspis kendallii (Gray): BMNH XXII.92a (lectotype, designated by Dring, 1979), Malaysia; FMNH 223201, MCZ 157158-59, Bako National Park, Sarawak, Malaysia; FMNH 223201; MCZ 157158-59, Bidi, Sarawak, Malaysia; FMNH 184424, Bukit Lanjan, Selangor, Malaysia; BMNH 1902.12.12.12, Bidi, Sarawak, Malaysia; BMNH 1911.1.20.7–9, Bau, Sarawak, Malaysia; BPBM 7494, Alag Sungei Ayer, Pulau Tioman, Pahang, Malaysia; ZRC 2.1101, Jerantut, Pahang, Malaysia; ZRC 2.1102, Gunung Rokan, Pulau Tioman, Pahang, Malaysia; ZRC 2.1103, Sedagong, Pulau Tioman, Pahang, Malaysia; ZRC 2.3014, Bukit Timah, Singapore; ZRC 2.3015, Gunung Ladang, Melaka, Malaysia; UF 78463, ZSI 14767, 19637 Malaysia; LSUHC 3773–75, 3797, 3811, 3820, 3841, 3878–88, 4659, 4666, 6213–15, 6218, 6224, Pulau Tioman, Pahang, Malaysia; LSUHC 3894, 5056–58, Pulau Tulai, Pahang, Malaysia; LSUHC 4707, 4756–57, 4765–67 Pulau Tinggi, Johor, Malaysia; LSUHC 4954, 4958 Sungai Lembing, Pahang, Malaysia; LSUHC 5184–87, 5198, 5211 Pulau Seribuat, Johor, Malaysia; LSUHC 5244 Pulau Sembilan, Johor, Malaysia; LSUHC 5523–24, 5731–34 Pulau Babi Besar, Johor, Malaysia; LSUHC 5532 Pulau Sibu, Johor, Malaysia; LSUHC 5703, 5711 Pulau Aceh, Pahang, Malaysia; LSUHC 5749–52 Pulau Babi Hunjung, Johor, Malaysia; LSUHC 6380–83 Pulau Ibol, Johor, Malaysia; LSUHC 6562 Kepong, Selangor, West Malaysia; LSUHC 7691, 8122, 8126, 8191, 8210, Endau-Rompin, Johor, Malaysia; ZRC 2.1109–10, Pulau Siantan, Anamba, Riao Archipelago, Indonesia; ZRC 2.1112–13, Sungei Ulu, Great Natuna, Riao Archipelago, Indonesia; USNM 26573, Pulau Bunoa, Tambelan Islands, Indonesia; USNM 26555, St. Barbe Island, Indonesia; USNM 26547–49, Bunguran, Natunas, Riao Archipelago, Indonesia; USNM 28145, Pulau Lingung, Natuna, Riao Archipelago, Indonesia; USNM 28149, Sirhassen, Natuna, Riao Archipelago, Indonesia, Cnemaspis kumpoli Taylor: LSUHC 8846–49, 8990–91, Wang Kelian, Perlis, Malaysia; MS 393–94, near the Ton Nga Chang Waterfall, Had Yai District, Songkhla Province (this is a new locality for this species). Cnemaspis limi Das & Grismer: ZRC 2.5289 (holotype), ZRC 2.3504–06, 2.5290 (paratypes), LSUHC 3801–02, 3859, 3902, 3904, 4410, 4425, 4480–83, 4485–88, 4563–64, 4596, 4604, 4616, 4629, 4655, 5053, 5424, 5441, 5510, 5515, 5518, 5521, 6203, 6206–07, 6210, 6212, 6267 Pulau Tioman, Pahang, Malaysia. Cnemaspis mcguirei Grismer, Grismer, Wood & Chan: LSUHC 8853–58, 8878, 8896, 8898, 9028–33, 9140, 9209, Bukit Larut, Perak, Malaysia; DWNP 1239, Gunung Bubu, Perak, Malaysia. Cnemaspis monachorum Grismer, Norhayati, Chan, Belabut, Muin, Wood, & Grismer: LSUHC 9113–19, Wat Wanararm, Kedah, Pulau Langkawi, Malaysia. Cnemaspis nigridius (Smith, 1925): BMNH 1946.8.22.90 (formerly BMNH 1925.9.1.8; holotype), MCZ 39024 and ZRC 2.1114-115, Mt. Gadin (= Gunung Gading), Sarawak, Malaysia; MCZ 15250, Lundu, Sarawak, Malaysia; BMNH 1925.9.1.9–10, Mt. Pueh, Sarawak, Malaysia. Cnemaspis nuicamensis Grismer & Ngo: LSUHC 9549–50, 9552–93, 9555, An Giang Prov., Tinh Bien Dist., Nui Cam Hill, Vietnam. Cnemaspis paripari Grismer & Chan: ZRC 2.6812, Gua Pari-pari, Bau District, Sarawak, Malaysia; LSUHC 9185, ZRC 2.6813–14, Gua Angin, Bau District, Sarawak, Malaysia. Cnemaspis pemanggilensis Grismer & Das: ZRC 2.6043 (holotype), ZRC 2.6044–51 (paratypes), LSUHC 4457–58, 4460, 4464, 4470–76, 4495–96, 8011–16, Pulau Pemanggil, Johor, Malaysia. Cnemaspis perhentianensis Grismer & Chan: LSUHC 8697, 8699, 9060, 9412, Pulau Perhentian Besar, Terengganu, Malaysia. Cnemaspis pseudomcguirei Grismer, Norhayati, Chan, Belabut, Muin, Wood, & Grismer: LSUHC 9047, 9145–47, Bukit Larut, Perak, Malaysia. Cnemaspis psychedelica Grismer, Ngo & Grismer: LSUHC 9182–85, Hon Khoai Island, Ngoc Hien District, Ca Mau Province, Vietnam. Cnemaspis roticanai Grismer & Chan: LSUHC 9430–31, 9439, 9453, Gunung Raya, Pulau Langkawi, Kedah, Malaysia. Cnemaspis tucdupensis Grismer & Ngo: LSUHC 8248, 9560–63, An Giang Prov., Tri Ton Dist., Tuc Dup Hill, Vietnam.

558 · Zootaxa 3691 (5) © 2013 Magnolia Press WOOD ET AL. 35 Chapter 3: Systematics and natural history of Southeast Asian Rock Geckos (genus

Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand,

and Indonesia

36 Zootaxa 3880 (1): 001–147 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3880.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:03A6448A-25D7-46AF-B8C6-CB150265D73D ZOOTAXA

3880

Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia

L. LEE GRISMER1,2, PERRY L. WOOD, JR3., SHAHRUL ANUAR4,5, AWAL RIYANTO6, NORHAYATI AHMAD2, MOHD A. MUIN4, MONTRI SUMONTHA7, JESSE L. GRISMER8, CHAN KIN ONN8, EVAN S. H. QUAH4 & OLIVIER S. A. PAUWELS9 1Department of Biology, La Sierra University, Riverside, California, USA E-mail: [email protected] 2Institute for Environment and Development, (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan, Malaysia. E-mail: [email protected] 3Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA, E-mail: [email protected] 4School of Biological Sciences, Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Penang, Malaysia, E-mail: [email protected], [email protected] 5Center for Marine and Coastal Studies, Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Penang, Malaysia, E-mail: [email protected] 6Museum Zoologicum Borgoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI), Cibinong, Bogor, Indonesia, E-mail: [email protected] 7Ranong Fisheries Marine Station, 157 M.1, Saphan-Pla Road, Paknam, Muang, Ranong 85000, Thailand, E-mail: [email protected] 8Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045-7651, USA, E-mail: [email protected], [email protected] 9Départmemt des Vertébrés Récens, Institut Royal des Sciences naturelles de Belgique, 29 Rue Vautier, 1000 Brussels, Belgium, E-mail: [email protected]

Magnolia Press Auckland, New Zealand

Accepted by A. Bauer: 29 Sept. 2014; published: 31 Oct. 2014 37 L. LEE GRISMER, PERRY L. WOOD, JR, SHAHRUL ANUAR, AWAL RIYANTO, NORHAYATI AHMAD, MOHD A. MUIN, MONTRI SUMONTHA, JESSE L. GRISMER, CHAN KIN ONN, EVAN S. H. QUAH & OLIVIER S. A. PAUWELS Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia (Zootaxa 3880) 147 pp.; 30 cm. 31 Oct. 2014 ISBN 978-1-77557-571-9 (paperback) ISBN 978-1-77557-572-6 (Online edition)

FIRST PUBLISHED IN 2014 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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2 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 38 Table of contents

Abstract ...... 4 Introduction ...... 4 Materials and methods ...... 6 Molecular analysis ...... 6 Phylogenetic analysis ...... 7 Morphological analysis ...... 7 Species accounts ...... 15 Results ...... 15 Taxonomy ...... 20 Ca Mau clade ...... 22 Cnemaspis boulengerii Strauch, 1887 ...... 23 Cnemaspis psychedelica Grismer, Ngo & Grismer, 2010 ...... 24 Pattani clade ...... 26 Cnemaspis monachorum Grismer, Norhayati, Chan, Belabut, Muin, Wood & Grismer, 2009 ...... 27 Cnemaspis biocellata Grismer, Chan, Nurolhuda & Sumontha, 2008 ...... 34 Cnemaspis niyomwanae Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010a ...... 36 Cnemaspis kumpoli Taylor, 1963 ...... 38 Northern Sunda clade ...... 39 Cnemaspis chanthaburiensis Bauer & Das, 1998 ...... 40 Cnemaspis neangthyi Grismer, Grismer & Thou, 2010 ...... 42 Cnemaspis aurantiacopes Grismer & Ngo, 2007 ...... 43 Cnemaspis caudanivea Grismer & Ngo, 2007 ...... 44 Cnemaspis nuicamensis Grismer & Ngo, 2007 ...... 46 Cnemaspis tucdupensis Grismer & Ngo, 2007 ...... 48 Cnemaspis siamensis (Smith, 1925) ...... 49 Cnemaspis huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 51 Cnemaspis chanardi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 52 Cnemaspis omari sp. nov...... 54 Cnemaspis roticanai Grismer & Chan, 2010 ...... 58 Cnemaspis flavigaster Chan & Grismer, 2008 ...... 60 Cnemaspis argus Dring, 1979 ...... 62 Cnemaspis karsticola Grismer, Grismer, Wood & Chan, 2008b ...... 63 Cnemaspis perhentianensis Grismer & Chan, 2008 ...... 64 Cnemaspis affinis (Stoliczka, 1870) ...... 66 Cnemaspis harimau Chan, Grismer, Shahrul, Quah, Muin, Savage, Grismer, Norhayati, Remegio & Greer, 2010b ...... 68 Cnemaspis pseudomcguirei Grismer, Norhayati, Chan, Belabut, Muin, Wood & Grismer, 2009 ...... 70 Cnemaspis shahruli Grismer, Chan, Quah, Mohd, Savage, Grismer,Norhayati, Greer & Remegio, 2010c ...... 72 Cnemaspis mcguirei Grismer, Grismer, Wood & Chan, 2008b ...... 74 Cnemaspis grismeri Wood, Quah, Anuar & Muin, 2013 ...... 79 Cnemaspis flavolineata (Nicholls, 1949) ...... 80 Cnemaspis temiah sp. nov ...... 81 Cnemaspis narathiwatensis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 85 Cnemaspis hangus sp. nov...... 86 Cnemaspis selamatkanmerapoh Grismer, Wood, Mohamed, Chan, Heinz, Sumarli, Chan & Loredo, 2013a ...... 90 Cnemaspis bayuensis Grismer, Grismer, Wood & Chan, 2008 ...... 90 Cnemaspis stongensis sp. nov...... 93 Southern Sunda clade ...... 97 Cnemaspis limi Das & Grismer, 2003 ...... 97 Cnemaspis nigridia (Smith, 1925) ...... 100 Cnemaspis paripari Grismer & Chan, 2009 ...... 100 Cnemaspis sundainsula sp. nov...... 103 Cnemaspis kendallii (Gray, 1845) ...... 107 Cnemaspis pemanggilensis Grismer & Das, 2006 ...... 111 Cnemaspis baueri Das & Grismer, 2003 ...... 112 Cnemaspis mumpuniae sp. nov...... 113 Cnemaspis peninsularis sp. nov...... 118 Cnemaspis bidongensis Grismer, Wood, Amirrudin, Sumarli, Vazquez, Ismail, Nance, Muhammad, Mohamad, Syed, Kuss, Murdoch & Cobos, 2014 ...... 124 Species Incertae Sedis ...... 126 Cnemaspis laoensis Grismer, 2010 ...... 126 Cnemaspis punctatonuchalis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 127 Cnemaspis vandeventeri Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 129

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 3 39 Cnemaspis kamolnorranathi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 ...... 130 Cnemaspis dringi Das & Bauer, 1998 ...... 131 Cnemaspis sundagekko sp. nov...... 134 Discussion ...... 137 Comments on biogeography ...... 137 Comments on diversity ...... 139 Comments on parallel evolution ...... 139 Comments on genetic divergence with Cnemaspsis ...... 141 Comments on integrative taxonomy ...... 141 Acknowledgements ...... 141 References ...... 141

Abstract

A well-supported and well-resolved phylogeny based on a concatenated data set from one mitochondrial and two nuclear genes, six morphological characters, and nine color pattern characters for 44 of the 50 species of the Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) is consistent with the previous taxonomy of Cnemaspis based solely on mor- phology and color pattern. Cnemaspis is partitioned into four major clades that collectively contain six species groups. The monophyly of all clades and species groups is strongly supported and they are parapatrically distributed across well- established, biogeographical regions ranging from southern Vietnam westward through southern Indochina, southward through the Thai-Malay Peninsula, then eastward to Borneo. Eight new species (Cnemaspis omari sp. nov. from the Thai- Malaysian border; C. temiah sp. nov. from Cameron Highlands, Pahang, Malaysia; C. stongensis sp. nov. from Gunung Stong, Kelantan, Malaysia; C. hangus sp. nov. from Bukit Hangus, Pahang, Malaysia; C. sundagekko sp. nov. from Pulau Siantan, Indonesia; C. peninsularis sp. nov. from southern Peninsular Malaysia and Singapore, and C. mumpuniae sp. nov. and C. sundainsula sp. nov. from Pulau Natuna Besar, Indonesia) are described based on morphology and color pattern and all but C. sundagekko sp. nov. are included in the phylogenetic analyses. Cnemaspis kendallii is polyphyletic and a composite of six species. An updated taxonomy consistent with the phylogeny is proposed for all 50 species and is based on 25 morphological and 53 color pattern characters scored across 594 specimens. Cladogenetic events and biogeograph- ical relationships within Cnemaspis were likely influenced by this group’s low vagility and the cyclical patterns of geo- graphical and environmental changes in Sundaland over the last 25 million years and especially within the last 2.5 million years. The phylogeny indicates that nocturnality, diurnality, substrate preferences, and the presence of ocelli in the shoul- der regions have evolved independently multiple times.

Key words: Cnemaspis, Gekkonidae, Southeast Asia, new species, Sunda Shelf, biogeography, phylogeography

Introduction

Southeast Asia harbors 20–25% of the planet’s terrestrial biodiversity in only 4% of its landmass. This makes it one of the great megadiverse hotspots of the world (Corlett 2009) even though much of its most prominent, geographic feature, the vast Sunda Plains, lie submerged beneath the South China Sea. Given that this is the second largest subareal margin of a continental shelf in the world (Parnell 2013) means that much of Sundaland’s terrestrial biodiversity is in a refugial state being restricted to small, Sundaic islands on the southern Sunda Plains and the areal, continental fringes of the Sunda Shelf (Grismer et al. 2011a). Furthermore, the current geographic outline of Sundaland has existed intermittently for only 2% of the last 2.4 million years (Woodruff 2010) owing to multiple, glaciostatic driven, sea-level changes that have repeatedly exposed and submerged the Sunda Plains while uniting Indochina with Borneo, Java, Sumatra, and the Thai-Malay Peninsula. Therefore, the phylogeographic relationships of Sundaic species and many species from southern Indochina cannot be accounted for by current geography and ecology alone but reconciled only in the context of cyclical changes in sea levels and concomitant climatic fluctuations (Cannon et al. 2009; Bell et al. 2010, 2011; Loredo et al. 2013; Woodruff 2010). One group that may offer insight into environmentally driven, cyclical speciation events in Sundaland is the monophyletic Southeast Asian branch of the polyphyletic gekkonid genus Cnemaspis Strauch (Gamble et al. 2012). Cnemaspis (sensu Smith 1933; Gamble et al. 2012) contains approximately 105 Afro-Asian scansorial species whose many morphological specializations are adaptations for moving about on flat, elevated surfaces during low levels of illumination. As such, the morphology within this genus appears to have been highly

4 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 40 conserved across the Old World. However, recent phylogenetic analyses (Gamble et al. 2012) have demonstrated that clades representing the major centers of radiation from Africa, South Asia, and Southeast Asia do not form a monophyletic group and are not particularly closely related to one another. Thus, their morphological similarity is due to convergence, not common ancestry. Working under this hypothesis, recent taxonomic revisions and descriptions of Southeast Asian Cnemaspis (Grismer et al. 2010a, 2013a, 2014; Wood et al. 2013) appropriately and expressly did not consider South Asian and African species in their analyses and only recognized Southeast Asian taxa related to the type species C. boulengerii Strauch, 1887 from southern Vietnam as true Cnemaspis.

FIGURE 1. Distribution of the Sundaland Rock Gecko genus Cnemaspis Strauch, 1887 in Southeast Asia. The northern outlier is C. laoensis Grismer 2009.

A monophyletic Cnemaspis sensu stricto as recognized throughout this study, currently contains 50 Sundaic and Indochinese species (see below) whose collective distribution encompasses the entire areal fringe of the Sunda Shelf and several islands across the submerged Sunda Plains (Fig. 1). The genus ranges from southern Laos in the north, southward through Vietnam, Cambodia, and Thailand, and south through the Thai-Malay Peninsula and

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 5 41 eastward to Borneo through the Seribuat, Anambas and Natuna archipelagos (Grismer & Chan 2010; Grismer et al. 2010a, 2013a, 2014; Wood et al. 2013). With some exceptions, all Cnemaspis are relatively small, cryptically colored species inhabiting primary and old secondary rainforests. They are secretive, microhabitat specialists that generally restrict their movements to the shaded surfaces of rocks, trees and caves during the day, or they are nocturnal. Having a body plan bearing a broad, flattened head; large, somewhat forward and upwardly directed eyes; a flattened body; and long, widely splayed limbs with long, inflected digits are adaptations for climbing on flat surfaces in all planes of orientation and show extreme interspecific conservatism despite a nearly three-fold difference in body size (maximum SVL ranging from 35.1 mm in C. monachorum to 88.2 mm in C. limi sp. nov.). This combination of cryptic behavior, microhabitat specialization, and morphological conservatism confounded species boundaries and generated considerable taxonomic confusion and instability (see Bauer & Das 1998; Chan & Grismer 2008; Das & Bauer 1998; Dring 1979; Grismer et al. 2008a,b, 2009). The confluence of these circumstances resulted in many species going unnoticed, unrecognized, or being buried in the synonymies of other species. However, a more informed understanding of how and where to look for Cnemaspis and what morphological characters are useful in establishing species boundaries has led to disentangling much of this group’s taxonomic history (see Grismer et al. 2010a and citations therein) along with the description of 43 new species since 2003. The morphological conservatism within Cnemaspis and the highly restricted distributions of its species coupled with their low vagility and microhabitat specialization, makes this genus particularly well-suited for comparative, biogeographic inferences in that multiple, parallel, cladogenetic events and phylogeographic patterns will have not have been erased through episodes of dispersal. The current taxonomy of Cnemaspis, however, is based solely on morphology and color pattern and in some cases, species were described using three or fewer specimens, potentially limiting the value of their diagnostic characters and the establishment of species boundaries. Furthermore, species boundaries and hypotheses of species relationships have never been tested with a molecular data set. Grismer et al. (2013b) demonstrated that morphology based taxonomies involving morphologically conserved taxa have great potential for underestimating species diversity, either due to cryptic speciation, character bias, or overlooked character states. They went on to provide evidence that the negative effects potentially inherent in such taxonomies can be significantly mitigated by an integrative approach (i.e., Padial et al. 2010). Therefore, the intent of this study is to test the morphological- based taxonomy of Cnemaspis using an integrative taxonomic approach to establish species level relationships in order to provide a platform for future analyses of this group’s potentially adaptive radiation and evolutionary biology (Wood & Grismer in prep.). We provide an expanded morphological and color pattern data set based on 596 specimens that include eight new and 43 previously described species in order to reinforce and modify their diagnoses and in some cases, significantly expand their geographic distributions and our knowledge of their natural history. All known species were included in the morphological and color pattern analyses and 44 of the 50 known species were included in the molecular analysis. These data are presented in a monographic style in order to consolidate information published in various books and journal articles over the last 130 years.

Materials and methods

Molecular analysis

A dataset from the mitochondrial gene NADH dehydrogenase subunit 2 gene (ND2) and its flanking tRNA’s (tRNAmet, tRNAtrp, tRNAala, tRNAasn, tRNAcys; 1335 bp), plus portions of two nuclear protein coding genes, Recombination Activating gene 1 (RAG-1; 1050 bp) and Matrix-Remodeling associated 5 gene (MXRA5; 839 bp) was constructed from 177 individuals of 44 of the 50 species of Cnemaspis from throughout its range plus 31 outgroups (Table 1) based on Gamble et al. (2012). Outgroup sequences were obtained from GenBank and all new sequences were deposited with GenBank accession numbers (Table 1). Two individuals per putative species (when available) were sequenced for the two nuclear genes. Previous studies have shown that using two samples per putative species can dramatically improve accuracy and precision (Camargo et al. 2012). Total genomic DNA was isolated from liver or skeletal muscle from specimens stored in 95% ethanol using the Qiagen DNeasyTM tissue kit (Valencia, CA, USA). All genes were amplified using a double-stranded Polymerase Chain Reaction (PCR) under the following conditions: 1.0 µl genomic DNA, 1.0 µl light strand primer 1.0 µl heavy strand primer, 1.0 µl dinucleotide pairs, 2.0 µl 5x buffer, 1.0 MgCl 10x buffer, 0.18 µl Taq polymerase, and 7.5 µl

6 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 42 H2O. PCR reactions were executed on an Eppendorf Mastercycler gradient theromocycler under the following conditions: initial denaturation at 95°C for 2 min, followed by a second denaturation at 95°C for 35 s, annealing at 47–52°C for 35 s, followed by a cycle extension at 72°C for 35 s, for 33–35 cycles (see Table 2 for details). All PCR products were visualized on a 10 % agarose gel electrophoresis. Successful targeted PCR products were vacuum purified using MANU 30 PCR plates Millipore plates and purified products were resuspended in DNAgrade water. Purified PCR products were sequenced using the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler. Cycle sequencing reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequenced on an ABI 3730xl DNA Analyzer at the BYU DNA Sequencing Center. Primers used for amplification and sequencing are presented in Table 2. All sequences were edited in Geneious v5.5.6 (Drummond et al. 2011) and initially aligned by eye. MacClade v4.08 (Maddison & Maddison 2005) was used to check for premature stop codons and to ensure the correct amino acid read frame for each gene.

Phylogenetic analysis

Both partitioned Maximum Likelihood (ML) and partitioned Bayesian Inference (BI) were employed. All genes were partitioned by codon position and the tRNA's were treated as one partition for both the ML and BI analyses. The Akaike Information Criterion (AIC) as implemented in ModelTest v3.7 (Posada & Crandall 1998), was used to calculate the best-fit model of evolution for each codon position (Table 3). Partitioned Maximum Likelihood analyses were performed using RAxML HPC v7.5.4 (Stamatakis, 2006), 1000 bootstrap pseudoreplicates via the rapid hill-climbing algorithm (Stamatakis et al. 2008) with 200 searches for the best tree. The Bayesian analysis was carried out in MrBayes v3.2 (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003) using the default priors. Two simultaneous runs were performed with eight chains per run, seven hot and one cold following default priors. The analysis was run for 2 x 106 generations and sampled every 1000 generations from the Markov Chain Monte Carlo (MCMC). The analysis was halted after the average standard deviation split frequency was below 0.01 and we assumed convergence. We conservatively discarded the first 25% of the trees as burnin. Nodes having bootstrap support values greater than 70 and posterior probabilities above 0.95 were considered significantly supported (Huelsenbeck et al. 2001; Wilcox et al. 2002). We calculated uncorrected percent sequence divergences for ND2 in Mega v5.2.2 (Tamura et al. 2011). Because it is possible for the mitochondrial and nuclear genes to have unique evolutionary histories (Hudson & Turelli 2003; Moore 1995), each gene was analyzed separately to look for discordances (Leaché & Mulcahy 2007). All analyses were performed using partitioned Maximum Likelihood implemented in RAxML HPC v7.5.4 (Stamatakis 2006) following the conditions described above and models of evolution in Table 3.

Morphological analysis

Color pattern characters were taken from digital images of living specimens cataloged in the La Sierra University Digital Photo Collection (LSUDPC) and from living specimens in the field. The following measurements on the type series were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate: snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90º to the inflection of the flexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90º to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eyeball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eyeball; eye to snout distance (ES), measured from anteriormost margin of the eyeball to the tip of snout; eye to nostril distance (EN), measured from the anteriormost margin of the eyeball to the

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 7 43 posterior margin of the external nares; inner orbital distance (IO), the width of the frontal bone at the level of the anterior edges of the orbit; ear length (EL), the greatest vertical distance of the ear opening; and internarial distance (IN), measured between the medial margins of the nares across the rostrum. Additional character states evaluated were numbers of supralabial and infralabial scales counted from below the middle of the orbit to the rostral and mental scales, respectively; the texture of the scales on the anterior margin of the forearm; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column (where applicable); the presence or absence of a row of enlarged, widely spaced, tubercles along the ventrolateral edge of the body (flank) between the limb insertions; the general size (i.e., strong, moderate, weak) and arrangement (i.e., random or linear) of the dorsal body tubercles; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the distribution of transverse and granular subdigital lamellae on the fourth toe; the total number of precloacal pores, their orientation and shape; the number of precloacal scales lacking pores separating the left and right series of pore-bearing precloacal scales; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales, subtibial scales, and submetatarsal scales beneath the first metatarsal; and the number of postcloacal tubercles on each side of the tail base. Longitudinal rows of caudal tubercles on the non-regenerated portion of the tail are quite variable between species and useful in differentiating several taxa. Up to five pairs of the following rows may be present in varying combinations: paravertebral row—the dorsal row adjacent to the middorsal, caudal furrow; dorsolateral row—the row between the paravertebral row and the lateral, caudal furrow on the dorsolateral margin of the tail; lateral row—the row immediately below the lateral, caudal furrow; and ventrolateral row—the row below the lateral row on the ventrolateral margin of the tail below the lateral caudal furrow. When present, this row is usually restricted to the anterior 25% (or less) of the tail. Occasionally there may be a row of tubercles within the lateral, caudal furrow. Fifty-three color pattern characteristics were evaluated and are described in Table 7. Much of the information on character states and their distribution throughout Cnemaspis was originally obtained from the literature and as such, was subject to inconsistencies between the methods of various authors in gathering and evaluating such data over the last 114 years. Grismer et al. (2010a), however, re-examined first-hand specimens of every species known at the time in order to standardize the character analysis. This is followed and augmented here with 25 morphological character states and 53 color pattern character states scored across all taxa (50 species totaling 596 specimens) as outlined above in the Materials and Methods. Additional material examined subsequent to the species’ original description or latest review is listed at the end of each species account. The distributions of the various species groups and clades constructed herein based on the molecular analysis closely correspond to particular biogeographic regions. This underscores the low vagility and microhabitat specificity of Cnemaspis species and provides a hypothetical framework for placing species that were not available for the molecular analysis in particular species groups and clades. The relationships of these species are left incertae sedis and their accounts are presented subsequent to the following proposed phylogenetic taxonomy. Institutional abbreviations follow Sabaj-Pérez (2010). ZRC refers to the Zoological Reference Collection at the Raffles Museum of Biodiversity, National University of Singapore, Singapore; DWNP refers to the Department of Wildlife and National Parks collection, Krau, Pahang, Peninsular Malaysia; LSUHC refers to the La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA; LSUDPC refers to the La Sierra University Digital Photo Collection; HC refers to the Herpetological Collection of the Universiti Kebangsaan Malaysia, Bangi, Selangor; USMHC refers to the Universiti Sains Malaysia Herpetological Collection at the Universiti Sains Malaysia, Penang, Malaysia; AMB refers to the collection of Aaron M. Bauer, Department of Biology, Villanova University, USA; THNHM refers to the Thailand Natural History Museum, National Science Museum, Khlong Luang, Pathum Thani, Bangkok, Thailand; CUMZ refers to the Chulalongkorn University Museum of Zoology, Bangkok, Thailand; PSUZC refers to the Prince of Songkhla University Zoological Collection, Songkhla, Thailand; UNS refers to the University of Natural Sciences in Ho Chi Minh City, Vietnam; ZMKU refers to the Zoological Museum of the Kasetsart University, Bangkok, Thailand; and KZM refers to the Nakhonratchasima Zoo Museum, ZPO, Nakhonratchasima, Thailand; CAS refers to the California Academy of Sciences, San Francisco, California, USA; FMNH refers to the Field Museum of Natural History, Chicago, Illinois, USA; MCZ refers to the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA; MNHN refers to the Museúm national d’Historie naturelle, Paris, France; MZB refers to the Museum Zoologicum Bogoriense, Cibinong, Indonesia; and TNHC refers to the Texas Natural heritage Collection and the University of Texas at Austin, Austin Texas, USA.

8 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 44 2 0 3 8 – – – – – – 6 2 U JN393959 JN393959 JQ945299 JQ945299 JQ945340 JQ945340 JN393986 JN393986 JX440679 JX440679 JN393984 JN393984 JX440678 JX440678 JX440676 EF534813 EF534813 EU054283 EU054283 EU268306 EU268306 E EU268299 HM559695 HM559695 HM559705 HM559705 HM997165 HM997165 – – – – XRA5 XRA5 RAG1 ……continued on the on next……continued page M alTissue Collection; AMS, Australian numbers accession GenBank al History; HC, Herpetological Collection of ense, Cibinung,Java, Indonesia; RAH, Rod – – – ND2 JX440515 JX440515 JX440566 JX440675 JN393914 JN393914 – JX041338 JX041338 – JN393943 JX440573 EF534785 JX440538 JX440594 JX440698 JX041448 JX041448 – JX440518 JX440572 EF534783 JN393948 JN393948 – JN393938 JN393938 – JX440517 JX440517 – JN393944 JN393944 – JX440516 JX440516 JX440567 JX440677 JX440520 JX440520 JX440576 JX440680 JX440519 JX440519 JX440574 EF534788 EU054299 EU054299 – EU268367 EU268367 – EU268363 EU268363 EU268360 – EU747728 EU747728 AAWZ02008741 AAWZ02015549 EU054288 EU054288 – AY369027 – GU459852 JX440577 GU459449 AY369025 – GQ257784 GQ257784 – GU459951 GU459951 JX440571 EF534779 HM559639 HM559639 – HM559629 HM559629 – HM997161 HM997161 – HM997153 HM997153 – Malaysia, Penang, Malaysia;USNM, United eum of Vertebrate Zoology (Berkeley); LSUHC, La La LSUHC, (Berkeley); Zoology Vertebrate eum of – Indicates missing data. data. missing – Indicates

y Localit USA: FloridaUSA: Myanmar:Rakhine State, Taung GokTownship Rajasthan, India,Rajasthan, Krom River Vietnam: An Giang Province, Tuc Dup Hill captive Tonga: Eua Island Australia, Queensland Australia Singapore USA: Louisiana Australia: Northern Territory Cambodia:Provinve, Pursat Keo Seima District Importedfrom Malaysia Indonesia:Krakatau Cambodia:Province, Pursat Phnom Aural unknown Ghana: Volta region, Togo Hills Myanmar:Tanintharyi Division, ThaungDistrict Kaw Thailand captive United Arab Emirates: Sharjah n/a captive Cuba Cuba USA: Puerto Rico Federated States ofMicronesia Australia: Northern Territory Pakistan:Balochistan, Makran district, Gwadar division Iran: Protected Area, Semnan Province New Zealand: Titahi Bay captive ecies p S Hemidactylus mabouia Hemidactylusmabouia Hemidactylusgarnotii Crossobamon orientalis Cyrtodactylus grismeri Mediodactylus russowii Mediodactylus Nactus pelagicus Oedura marmorata Lialis jicari Lepidodactylus lugubris lugubris Lepidodactylus Hemidactylusturcicus Heteronotia binoei Dixonius siamensis fehlmanni Gehyra Gehyra insulensis mutilata Gehyra geckoGekko Hemidactylusangulatus Hemidactylusfrenatus Dixonius melanostictus Cyrtopodion elongatum Bunopus tuberculatus Bunopus Anolis carolinensis Stenodactylus petriiStenodactylus Sphaerodactylus torreiSphaerodactylus Sphaerodactylus rooseveltiSphaerodactylus Perochirusateles nigriceps Pygopus Agamura persica Tenuidactylus caspius Woodworthia maculata maculata Woodworthia Tropicolotes steudneri Specimensfor used the molecularphylogenetic analyses. Vouchernumber abbreviations are asfollows: AustralianABTC, Biologic Voucher numberVoucher M 14798 14798 M YP CAS 223286 223286 CAS ID 7618 ID7618 LSUHC 8638 8638 LSUHC n/a 322160 USNM 143861 AMS n/a ZRC 24847 24847 ZRC LSUMZ H-1981 LSUMZ H-1981 R6147 ANWC LSUHC 7328 7328 LSUHC 00723 TG 13940 ABTC 7379 LSUHC n/a MVZ245438 229633 CAS n/a JB 127 127 JB CAS 228737 228737 CAS States NationalMuseum (Smithsonian);Yale YPM, PeabodyMuseum; ZRC, ZoologicalReference Collection,Raffles Museum. n/a TABLE 1. TABLE Museum,Sydney; Australian ANWC, National Wildlife Collection;CaliforniaCAS, Academyof Sciences; FMNH, Field Museum of Natur the Universiti KebangsaanMalaysia, Bangi,Selangor; ID, IndraneilDas field series; Boone Jon JB, captive collection; MVZ,Mus A. Hitchmoughfield series; Gamble; TG, Tony Universiti USMHC, Malaysia Sains Herpetological Collection atthe Universiti Sains Sierra University Herpetological Collection;LSUMZ,Louisiana State University Museum of Zoology; MZB, Museum Zoologicum Bogori JB 35 35 JB JB 34 34 JB CAS 198428 198428 CAS n/a MVZ197233 FMNH 274474 274474 FMNH CAS 228602 228602 CAS RAH292 RAH292 JB 28 28 JB

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 9 45

8 2 9 4

2 – – – – – – – – – – – – – – – – – – – – – – – – – – 0

M KM024930 K

2 6 6 5 8 8 4 4

2 2 – – – – – – – – – – 024858 0 0 ……continued on the on next……continued page M M KM KM024857 K K

7 7 9 6 4 GenBank accession numbers accession GenBank 8 9 9 9 8 6 6 6 6 6 4 4 4 4 4 2 2 2 2 2 0 0 0 0 0 ND2 ND2 MXRA5 RAG1 M M M M M KM024694 KM024694 KM024861 KM024931 KM024693 KM024693 – KM024710 KM024710 KM024866 KM024692 KM024692 KM024860 KM024711 KM024711 KM024867 KM024716 KM024716 KM024871 KM024688 KM024688 KM024689 – KM024695 KM024695 – KM024714 KM024714 K KM024709 KM024709 KM024865 KM024934 KM024707 KM024707 KM024708 KM024864 KM024933 – KM024698 KM024698 – KM024712 KM024713 KM024868 KM024869 KM024935 KM024936 KM024686 KM024686 KM024681 KM024681 KM024701 KM024701 KM024706 KM024682 KM024682 – KM024700 – KM024683 KM024683 KM024705 – K K K K KM024704 KM024704 KM024702 KM024702 KM024863 KM024932 KM024685 KM024703 KM024703 – KM024690 KM024690 KM024691 KM024859 – KM024929 t i w a

L

g g Locality Locality n u n r r r u G

, u n a g g n e r e T

: a i s y a l a Malaysia: Terengganu, GunungTebu Malaysia: Terengganu, GunungTebu Malaysia: Terengganu, Pulau Bidong M Malaysia: Penang, Pulau Pinang Malaysia: Penang, Pulau Pinang Malaysia: Penang, Pulau Pinan Malaysia: Kelantan,Kampung Bayu Malaysia: Perlis, Kuala Perlis Vietnam: CaMauProvince, Con DaoArchipelago Malaysia: Kelantan,Kampung Bayu Malaysia: Terengganu, Pulau Bidong Malaysia: Perlis, GuaKelam Malaysia: Perlis, Kuala Perlis SiThailand: Nakhon Thammarat Province, Thum Thong Panra Cambodia:Province, Pursat Phnom Dalai KM024715 KM024870 KM024937 Malaysia: Penang, Pulau Pinang Malaysia: Penang, Pulau Pinang Malaysia: Terengganu, GunungTebu Malaysia: Johor,PulauAu Malaysia: Kelantan,Kampung Bayu Malaysia: Terengganu, GunungTebu Malaysia: Johor,PulauAu Vietnam: Kien GiangProvince, HonDat Hill Malaysia: Johor,PulauAur Vietnam: Kien Giang Province, Hon Tre Island Vietnam: Kien GiangProvince, HonTre Island Malaysia: Johor,PulauAur Vietnam: Kien Giang Province, Hon Tre Island Vietnam: Kien GiangProvince, HonDat Hill Malaysia: Terengganu, Pulau Bidong Vietnam: CaMauProvince, Con DaoArchipelago Malaysia: Terengganu, Pulau Bidong Malaysia: Penang, Pulau Pinang Vietnam: Kien GiangProvince, HonDat Hill Malaysia: Johor,PulauAu

inis Species ff is a is p Cnemaspis argus Cnemaspis Cnemaspis argus Cnemaspis bidongensis Cnemaspis affinisCnemaspis Cnemaspis aurantiacopes Cnemas Cnemaspis bayuensis biocellata Cnemaspis Cnemaspis boulengerii Cnemaspis chanthaburiensis Cnemaspis bayuensis Cnemaspis bidongensis biocellata Cnemaspis biolcellataCnemaspis Cnemaspis chanardi Cnemaspis baueri Cnemaspis Cnemaspis bayuensis Cnemaspis affinisCnemaspis affinisCnemaspis Cnemaspis argus Cnemaspis argus Cnemaspis argus Cnemaspis affinisCnemaspis Cnemaspis baueri Cnemaspis Cnemaspis baueri Cnemaspis Cnemaspis caudanivea caudanivea Cnemaspis caudanivea Cnemaspis Cnemaspis baueri Cnemaspis caudanivea Cnemaspis Cnemaspis aurantiacopes Cnemaspis bidongensis boulengerii Cnemaspis Cnemaspis bidongensis Cnemaspis affinisCnemaspis Cnemaspis aurantiacopes Cnemaspis baueri Cnemaspis

Voucher numberVoucher LSUHC 10834 10834 LSUHC 10835 LSUHC LSUHC 11447 11447 LSUHC TABLE 1. (Continued) 1. TABLE 6774 LSUHC LSUHC 6757 6757 LSUHC LSUHC 9071 9071 LSUHC 8818 LSUHC 9278 LSUHC LSUHC 9338 9338 LSUHC LSUHC 8817 8817 LSUHC 9567 LSUHC LSUHC 9072 9072 LSUHC 11446 LSUHC 8789 LSUHC LSUHC 7303 7303 LSUHC 9070 LSUHC LSUHC 6758 LSUHC 8304 LSUHC 10858 LSUHC LSUHC 6787 6787 LSUHC LSUHC 10859 10859 LSUHC LSUHC 6759 6759 LSUHC LSUHC 7302 7302 LSUHC LSUHC 7301 7301 LSUHC LSUHC 8578 8578 LSUHC 8582 LSUHC LSUHC 8612 8612 LSUHC 7273 LSUHC 8577 LSUHC LSUHC 8611 8611 LSUHC LSUHC 11445 11445 LSUHC 9279 LSUHC LSUHC 11444 11444 LSUHC LSUHC 6788 6788 LSUHC LSUHC 8610 8610 LSUHC LSUHC 7274 7274 LSUHC

10 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 46

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2 – – – – – – – – – – – – – – – – – – – – 0

M KM024940 K 9 7 8 4

2 – – – – – – – – – 0 ……continued on the on next……continued page M K

3 GenBank accession numbers accession GenBank 3 7 4 2 0 ND2 MXRA5 RAG1 M KM024734 KM024734 – KM024735 KM024735 KM024880 KM024944 KM024745 KM024745 KM024746 KM024885 KM024886 KM024950 KM024949 KM024720 KM024720 KM024872 KM024738 KM024738 KM024736 KM024736 KM024737 KM024881 KM024882 KM024945 KM024946 KM024721 KM024721 KM024874 KM024938 KM024742 KM024742 KM024741 KM024741 – KM024739 KM024739 – KM024740 KM024740 KM024884 KM024743 KM024948 KM024883 KM024947 KM024751 – KM024718 KM024718 KM024873 KM024744 KM024744 – KM024749 KM024752 – KM024889 KM024953 KM024726 KM024726 – KM024724 KM024724 KM024725 – KM024719 KM024719 KM024722 – KM024875 KM024939 KM024748 KM024748 KM024727 KM024727 KM024728 KM024729 KM024876 KM024941 KM024730 KM024730 KM024731 KM024732 KM024877 KM024878 KM024942 KM024717 KM024717 K KM024723 KM024747 KM024747 KM024888 KM024952 KM024750 KM024750 KM024887 KM024951 National Park Locality Locality

g Thailand: Kanchanaburi Province, Sai Yok National Park Thailand: Kanchanaburi Province, Sai Yok National Park Malaysia: Kelantan,RengGunung Malaysia: Kelantan,RengGunung Indonesia:Riau Province, Pulau Serasan Malaysia: Sarawak, Gunung Gading Malaysia: Selangor,Kepong Malaysia: Selangor,Ulu Gombak Malaysia: Sarawak, Gunung Gading Malaysia:Jerai Kedah,Gunung Malaysia: Sarawak, Gunung Gading Malaysia: Pahang, Pulau Tioman Malaysia: Perak, Lenggong Malaysia: Pahang, Bukit Hangus Malaysia: Pahang, Bukit Hangus Malaysia: Pahang, Bukit Hangus Malaysia:Jerai Kedah,Gunung Malaysia:Jerai Kedah,Gunung Malaysia: Sarawak, Gunung Gading Malaysia: Sarawak, Santubong Malaysia: Selangor,Kepon Malaysia: Perak, Lenggong Malaysia: Perak, Lenggong Malaysia: Perak, Lenggong Malaysia: Perak, Lenggong Malaysia: Selangor,Kepong Malaysia: Pahang, Fraser’s Hill, The Gap Thailand: Kanchanaburi Province, Sai Yok Malaysia: Perlis, Perlis StatePark Malaysia: Pahang, Pulau Tioman Malaysia: Pahang, Pulau Tioman Malaysia: Perlis, Perlis StatePark Malaysia: Perak, BukitLarut Malaysia: Sarawak, Santubong Malaysia: Pahang, Pulau Tioman Malaysia: Perak, BukitLarut

sp.nov. sp.nov. sp.nov. aster g Species lavi f is is p Cnemaspis huaseesom huaseesom Cnemaspis Cnemaspis huaseesom karsticolaCnemaspis Cnemaspis karsticolaCnemaspis kendallii Cnemaspis Cnemaspis kendallii Cnemaspis Cnemaspis flavigaster Cnemas Cnemaspis flavigasterCnemaspis Cnemaspis kendallii Cnemaspis Cnemaspis harimau kendallii Cnemaspis Cnemaspislimi Cnemaspis kendallii Cnemaspis kendallii Cnemaspis Cnemaspis grismeriCnemaspis Cnemaspis hangus Cnemaspis hangus Cnemaspis hangus Cnemaspis harimau Cnemaspis harimau Cnemaspis grismeri grismeriCnemaspis Cnemaspis grismeri grismeriCnemaspis Cnemaspis flavigasterCnemaspis flavolineata Cnemaspis Cnemaspis huaseesom Cnemaspis kumpoli Cnemaspis Cnemaspis limi Cnemaspis limi Cnemaspis kumpoli Cnemaspis mcguireiCnemaspis Cnemaspis kendallii Cnemaspis limiCnemaspis Cnemaspis mcguirei

Voucher numberVoucher LSUHC 9457 9457 LSUHC 9458 LSUHC 9054 LSUHC LSUHC 9055 9055 LSUHC 5317 LSUHC LSUHC 9172 9172 LSUHC TABLE 1. (Continued) 1. TABLE 8835 LSUHC LSUHC 6562 6562 LSUHC LSUHC 10380 10380 LSUHC LSUHC 9174 9174 LSUHC LSUHC 9668 9668 LSUHC 9173 LSUHC LSUHC 3774 3774 LSUHC LSUHC 9171 9171 LSUHC 9178 LSUHC HC 0225 0225 HC 9667 LSUHC LSUHC 9733 9733 LSUHC 9358a LSUHC 9358b LSUHC 9665 LSUHC LSUHC 9730 9730 LSUHC 9732 LSUHC LSUHC 9969 9969 LSUHC 9970 LSUHC LSUHC 8836 8836 LSUHC 8079 LSUHC 9455 LSUHC LSUHC 8848 8848 LSUHC 3902 LSUHC 3904 LSUHC LSUHC 8847 8847 LSUHC LSUHC 8853 8853 LSUHC LSUHC 9179 9179 LSUHC 3888 LSUHC LSUHC 8854 8854 LSUHC

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M KM024957 K KM024969 KM024969

ontinued on the on next ontinued page

– – – – – – – – – – – – ……c KM024894

1 GenBank accession numbers accession GenBank 8 7 4 2 0 ND2 ND2 MXRA5 RAG1 M KM024776 KM024902 KM024778 KM024778 KM024903 KM024965 KM024775 – KM024777 KM024779 – KM024904 KM024966 KM024761 KM024782 KM024782 KM024783 KM024905 KM024784 KM024906 KM024967 KM024968 KM024768 KM024768 KM024753 KM024753 KM024890 KM024954 KM024766 KM024766 KM024767 KM024767 KM024897 KM024769 KM024898 KM024960 KM024961 KM024764 KM024764 KM024765 KM024896 KM024959 KM024771 KM024762 KM024762 KM024895 KM024958 KM024755 KM024755 KM024758 – KM024786 KM024786 KM024788 KM024785 KM024785 KM024787 – – KM024763 KM024763 KM024770 KM024899 KM024772 KM024900 KM024963 KM024962 KM024773 KM024773 KM024774 KM024901 KM024964 K KM024780 KM024780 – KM024759 KM024756 KM024756 KM024757 KM024892 KM024955 – KM024760 KM024760 KM024893 KM024956

ari Locality Locality Vietnam: An Giang Province, Nui Cam Hill Malaysia: Perak, BukitLarut Malaysia: Kedah,Langkawi Archipelago, Pulau Langkawi KM024754 KM024891 Malaysia: Sarawak, Gua Pari-P Vietnam:Giang An Province, CamNui Hill Malaysia: Perlis, Perlis StatePark Thailand: Satun Province, Cave Phuphaphet Malaysia: Johor,PulauPemanggil Malaysia: Johor,PulauPemanggil Malaysia: Sarawak, Gua Angin Malaysia: Sarawak, Gua Angin Malaysia: Sarawak, Gua Angin Malaysia: Johor,PulauPemanggil Malaysia: Johor,PulauPemanggil Indonesia:Riau Province, Bedung fragmentedforest area Malaysia: Perak, Belum-Temengor, Sungai Enam Malaysia: Perak, Belum-Temengor, Sungai Enam Cambodia: PursatProvince, O’Lakmeas Cambodia:Province, Pursat O’Lakmeas Cambodia: PursatProvince, O’Lakmeas Malaysia: Sarawak, Gunung Gading Malaysia: Sarawak, Gunung Gading Vietnam: An Giang Province, Nui Cam Hill Malaysia: Perak, Belum-Temengor, Sungai Enam Malaysia: Kedah,Langkawi Archipelago, Pulau Langgun Vietnam: An Giang Province, Nui Cam Hill Malaysia: Kedah,Langkawi Archipelago, Pulau Langgun Malaysia: Kedah,Langkawi Archipelago, Pulau Langgun Malaysia: Kedah,Langkawi Archipelago, Pulau Langgun Cambodia:Province, Pursat O’Lakmeas Thailand: Trang Province,Thum Ting Khao Malaysia: Sarawak, Gunung Gading Thailand: Trang Province,Thum Ting Khao Malaysia: Perak, Belum-Temengor, Sungai Enam

sp. nov. Indonesia:Riau Province, Sekunyam ForestReserve sp. nov. sp. nov. sp. Reserve Forest Sekunyam Province, Riau Indonesia:

sp. nov.sp. nov.sp. uirei g Species is mc is p Cnemaspis nuicamensis Cnemaspis monachorum Cnemas Cnemaspis paripari Cnemaspis nuicamensis omari Cnemaspis omari Cnemaspis Cnemaspis pemanggilensisCnemaspis pemanggilensisCnemaspis Cnemaspis mumpuniae Cnemaspis paripari Cnemaspis paripari Cnemaspis paripari Cnemaspis pemanggilensis Cnemaspis pemanggilensis Cnemaspis nuicamensis Cnemaspis mumpuniae mumpuniae Cnemaspis Cnemaspis narathiwatensis Cnemaspis narathiwatensis Cnemaspis neangthyi Cnemaspis neangthyi Cnemaspis neangthyi Cnemaspis nigridia Cnemaspis nigridia Cnemaspis narathiwatensis Cnemaspis monachorum Cnemaspis nuicamensis Cnemaspis monachorum monachorum Cnemaspis Cnemaspis monachorum Cnemaspis monachorum Cnemaspis neangthyi Cnemaspis niyomwanae Cnemaspis nigridia Cnemaspis niyomwanae Cnemaspis narathiwatensis narathiwatensis Cnemaspis

Voucher numberVoucher LSUHC 8647 8647 LSUHC TABLE 1. (Continued) 1. TABLE 9114 LSUHC LSUHC 8855 8855 LSUHC LSUHC 9184 9184 LSUHC LSUHC 8649 8649 LSUHC 9978 LSUHC 9565 LSUHC LSUHC 8012 8012 LSUHC 8014 LSUHC MZB.Lace 10167 10167 MZB.Lace 9186 LSUHC 9192 LSUHC LSUHC 9185 9185 LSUHC 8011 LSUHC 8013 LSUHC LSUHC 8646 8646 LSUHC USMHC 1350 1350 USMHC 8515 LSUHC 8517 LSUHC 9170 LSUHC MZB.Lace 10155 MZB.Lace 10166 MZB.Lace 1349 USMHC 8516 LSUHC 9168 LSUHC USMHC 1347 1347 USMHC LSUHC 10808 10808 LSUHC 8648 LSUHC LSUHC 10809 10809 LSUHC LSUHC 10807 10807 LSUHC 10810 LSUHC LSUHC 8478 8478 LSUHC 9571 LSUHC LSUHC 9169 9169 LSUHC 9568 LSUHC USMHC 1348 1348 USMHC

12 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 48

– – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – –

– – – – – – – – – – – – – – – – – – – ……continued on the on next ……continued page

GenBank accession numbers accession GenBank ND2 ND2 MXRA5 RAG1 KM024797 KM024797 KM024795 KM024795 KM024799 KM024800 KM024802 KM024803 – KM024814 KM024814 KM024815 KM024816 – KM024824 KM024824 KM024910 KM024973 KM024792 KM024792 KM024793 KM024796 – – KM024801 – KM024806 – KM024790 KM024790 KM024791 – KM024798 – KM024805 KM024907 KM024789 KM024789 KM024817 KM024817 KM024818 KM024819 – KM024794 KM024794 KM024804 KM024807 KM024808 KM024809 – KM024821 KM024821 KM024909 KM024823 KM024972 – KM024810 KM024810 KM024811 KM024908 KM024812 KM024813 KM024971 – – KM024820 KM024820 KM024822

Locality Locality Malaysia: Pahang, Pulau Tioman Malaysia: Pahang, Pulau Tioman Malaysia: Johor,PulauTinggi Malaysia: Johor,PulauTinggi Malaysia: Pahang, Sungai Lembing Malaysia: Johor,PulauTulai Malaysia: Johor,PulauSeribuat Malaysia: Johor,PulauSeribuat Malaysia: Johor,PulauAceh Malaysia: Johor,PulauAceh Malaysia: Johor,PulauIbol Malaysia: Johor,PulauIbol Malaysia: Johor,PulauBabi Besar Malaysia: Johor,PulauBabi Besar Malaysia: Johor,Endau-Rompin, Selai Malaysia: Johor,Endau-Rompin, Selai Malaysia:Panti Johor,Gunung Forest Reseve Malaysia:Ledang Johor,Gunung Malaysia:Ledang Johor,Gunung Malaysia:Ledang Johor,Gunung Malaysia: Terengganu, Pulau Tenggol Malaysia: Terengganu, Pulau Tenggol Malaysia: Terengganu, Pulau Tenggol Malaysia: Terengganu, Pulau Tenggol Malaysia:Berlumut Johor,Gunung Malaysia:Berlumut Johor,Gunung Singapore: NeeSwampSoon Malaysia: Pahang, Bukit Hangus Malaysia: Pahang, Bukit Hangus Malaysia: Pahang, Bukit Hangus Malaysia: Terengganu, Pulau Perhentian Besar Malaysia: Terengganu, Pulau Perhentian Besar Malaysia: Terengganu, Pulau Perhentian Besar Malaysia: Terengganu, Pulau Perhentian Besar Malaysia: Perak, BukitLarut Malaysia: Pahang, Pulau Tioman Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó Ó

sp. nov.sp. nov.sp. sp. nov.sp. sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. sp. nov.sp. sp. nov.sp. sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. nov.sp. sp. nov.sp. sp. nov.sp. nov.sp. sp. nov.sp. nov.sp. Species eninsularis p is is p Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemas Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis perhentianensis Cnemaspis peninsularis Cnemaspis perhentianensis Cnemaspis perhentianensis Cnemaspis perhentianensis Cnemaspis pseudomcguirei Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis Cnemaspis peninsularis

Voucher numberVoucher LSUHC 4958 4958 LSUHC 8965 LSUHC TABLE 1. (Continued) 1. TABLE 3797 LSUHC LSUHC 3773 3773 LSUHC LSUHC 5703 5703 LSUHC 5523 LSUHC 8122 LSUHC 9380 LSUHC 10238 LSUHC 0226 HC LSUHC 9377 9377 LSUHC LSUHC 9060 9060 LSUHC LSUHC 9145 9145 LSUHC HC 0228A 0228A HC 8699 LSUHC 8700 LSUHC 9412 LSUHC LSUHC 9376 9376 LSUHC 9382 LSUHC 10239 LSUHC 10454 LSUHC LSUHC 6381 6381 LSUHC 5731 LSUHC 8910 LSUHC 8966 LSUHC LSUHC 4756 4756 LSUHC 5056 LSUHC 5184 LSUHC 5307 LSUHC 6380 LSUHC LSUHC 3820 3820 LSUHC LSUHC 5185 5185 LSUHC 0228B HC LSUHC 4707 4707 LSUHC 8967 LSUHC LSUHC 8126 LSUHC

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 13 49

– – – – – – – – – – – – – – – – – –

KM024978 KM024978 KM024985 KM024985 KM024976

– – – – – – – – KM024917 KM024915

GenBank accession numbers accession GenBank – ND2 ND2 MXRA5 RAG1 043K041 KM024977 M024833KM024916 KM024826 KM024826 KM024852 KM024852 KM024926 KM024855 KM024986 KM024927 KM024987 KM024853 KM024854 – KM024827 KM024827 KM024912 KM024825 KM024825 KM024911 KM024974 KM024828 KM024828 KM024837 – KM024829 KM024829 KM024830 KM024913 – KM024849 KM024849 KM024850 KM024925 KM024984 KM024851 KM024851 – KM024835 KM024835 KM024918 KM024840 KM024840 KM024841 KM024921 KM024842 KM024922 KM024980 – KM024981 KM024831 KM024831 KM024914 KM024975 KM024836 K KM024834 KM024843 KM024843 – KM024839 KM024839 KM024920 KM024838 KM024838 KM024919 KM024979 KM024845 KM024845 KM024846 KM024923 KM024847 KM024924 KM024982 KM024983 KM024832

Locality Locality Malaysia: Pahang, Merapoh, Gua Gunting Gua Merapoh, Pahang, Malaysia: Malaysia: Kedah,PulauLangkawi, Raya Gunung Malaysia: Kedah,PulauPinang Malaysia: Perak, BukitLarut Malaysia: Pahang, Cameron Highlands,Rata Tanah Malaysia: Pahang, Cameron Highlands,Rata Tanah Vietnam: An Giang Province, Tuc Dup Hill Indonesia:Riau Province, Besar, Natuna Pulau CerukForest Indonesia:Riau Province, Besar, Natuna Pulau CerukForest Malaysia: Pahang, Cameron Highlands,Rata Tanah Malaysia: Penang, Pulau Jerejak Indonesia:Riau Province, Besar, Natuna Pulau CerukForest Indonesia:Riau Province,Besar, Pulau Natuna Air Gunung Hiu KM024848 Vietnam:Giang An Province, Hill Tuc Dup – Vietnam:Giang An Province, Hill Tuc Dup Vietnam:Mau Can Province, HonKhoai Island Malaysia: Kedah,PulauLangkawi, Raya Gunung Malaysia: Perak, BukitLarut Malaysia: Penang, Pulau Pinang Malaysia: Kelantan,Stong,Gunung Kem Baha Malaysia: Kelantan,Stong,Gunung Kem Baha Thailand: Chumpon Province,Thailand: Chumpon Pathio Malaysia: Kedah,PulauLangkawi, Raya Gunung Malaysia: Kedah,Sungai Sedim Province,Thailand: Chumpon Pathio Malaysia: Pahang, Merapoh,Gua Gunting Vietnam: An Giang Province, Tuc Dup Hill Vietnam:Mau Can Province, HonKhoai Island Malaysia: Penang, Pulau Jerejak Indonesia:Riau Province,Besar Pulau Natuna Malaysia: Kelantan,Stong,Gunung Kem Baha Indonesia:Riau Province,Besar, Pulau Natuna Ranai Gunung KM024844

sp. nov.sp. nov.sp. sp. nov.sp. nov.sp. sp. nov.sp. sp. nov.sp. uirei sp. nov. sp. g

sp. nov. sp. sp. nov. sp. sp. nov. sp. Species seudomc p is is p Cnemaspis roticanai Cnemaspis shahruli Cnemaspis pseudomcguirei Cnemas Cnemaspis temiah Cnemaspis temiah Cnemaspis tucdupensis Cnemaspis sundainsula Cnemaspis Cnemaspis sundainsula Cnemaspis shahruli Cnemaspis sundainsula Cnemaspis sundainsula Cnemaspis temiah Cnemaspis tucdupensis Cnemaspis tucdupensis Cnemaspis psychedelicaCnemaspis Cnemaspis roticanai Cnemaspis shahruli Cnemaspis stongensisnov. sp. Cnemaspis stongensisnov. sp. Cnemaspis siamensis Cnemaspis roticanai Cnemaspis shahruli Cnemaspis siamensis Cnemaspis selamatkanmerapoh Cnemaspis Cnemaspis tucdupensis Cnemaspis psychedelicaCnemaspis Cnemaspis shahruli Cnemaspis stongensis sundainsula Cnemaspis Cnemaspis sundainsula Cnemaspis selamatkanmerapoh

Voucher numberVoucher LSUHC 9431 9431 LSUHC 10375 LSUHC LSUHC 9147 9147 LSUHC TABLE 1. Continued. 1. TABLE LSUHC 9146 9146 LSUHC LSUHC 9739 9739 LSUHC LSUHC 9160 9160 LSUHC 8633 LSUHC MZB.Lace 10157 10157 MZB.Lace 10158 MZB.Lace LSUHC 9816 9816 LSUHC 8634 LSUHC MZB.Lace 10156 10156 MZB.Lace 10160 MZB.Lace 8631 LSUHC LSUHC 9613 9613 LSUHC LSUHC 9243 9243 LSUHC 9430 LSUHC LSUHC 6773 6773 LSUHC LSUHC 11091 11091 LSUHC LSUHC 11089 11089 LSUHC LSUHC 9485 9485 LSUHC LSUHC 9439 9439 LSUHC 9586 LSUHC 9474 LSUHC LSUHC 11015 LSUHC LSUHC 8632 8632 LSUHC LSUHC 9244 9244 LSUHC LSUHC 9163 9163 LSUHC LSUHC 11090 11090 LSUHC 5314 LSUHC MZB.Lace 9436 9436 MZB.Lace LSUHC 11016 11016 LSUHC

14 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 50 TABLE 2. A list of primers used for amplification and sequencing. (int)=internal used for sequencing only; cyc/temp= number of cycles/annealing temperature.

Gene Primer name Primer reference Sequence cyc/temp ND2 L4437b (Macey & Schulte, 1999) 5’-AAGCAGTTGGGCCCATACC-3’ 33/48 L5002 (Macey & Schulte, 1999) 5’-AACCAAACCCAACTACGAAAAAT-3’ CYRTINTF1(int) (Siler et al., 2010) 5’-TAGCCYTCTCYTCYATYGCCC-3’ CYRTINTR1(int) (Siler et al., 2010) 5’-ATTGTKAGDGTRGCYAGGSTKGG-3’ MXRA5 MXRA5F2 (Portik et al., 2011) 5’-KGCTGAGCCTKCCTGGGTGA-3’ 35/66 MXRA5R2 (Portik et al., 2011) 5’-YCTMCGGCCYTCTGCAACATTK-3’ RAG-1 R13 (Groth & Barrowclough, 1999) 5’-TCTGAATGGAAATTCAAGCTGTT-3’ 35/56 R18 (Groth & Barrowclough, 1999) 5’-GATGCTGCCTCGGTCGGCCACCTTT-3’

TABLE 3. Selected and applied models of molecular evolution for data partitions determined by AIC for the phyloge- netic analyses.

Gene Models selected Models applied ND2 1st pos GTR+I+Γ GTR+I+Γ 2nd pos GTR+I+Γ GTR+I+Γ 3rd pos GTR+I+Γ GTR+I+Γ tRNAs TrN+I+ Γ HKY+I+Γ MXRA5 1st pos HKY+Γ HKY+Γ 2nd pos TIM+I HKY+Γ 3rd pos K81uf+ Γ GTR+I RAG-1 1st pos K81uf+ Γ GTR+ Γ 2nd pos K81uf+ Γ GTR+ Γ 3rd pos TVM+ Γ GTR+ Γ

Species accounts

Accounts for each species are presented in phylogenetic order based on Figure 2 from most basal to most derived. Accounts of previously described species are bolstered (in most cases) with data from additional specimens and observations. All accounts include information on the holotype and type locality, a diagnosis, a color pattern description, distribution, natural history (which includes previously published data modified and supplemented by additional observations), phylogenetic relationships, remarks (if necessary), and material examined subsequent to the original description. A synonymy of all names used is presented but only major references or commonly cited references are listed. If the account is a new species description, sections on the holotype, paratypes, variation, comparisons to other species, and etymology are included as well.

Results

Data from all three loci (ND2, MXRA5, and RAG1) were analyzed separately and the resulting trees were compared. ND2 was far superior at resolving relationships in the more shallow regions of the tree whereas these relationships were largely unresolved by the nuclear genes. However, ND2 was unable to resolve some of the basal

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 15 51 relationships in deeper regions of the tree where both nuclear genes produced the same phylogenetic sequence among basal nodes.

FIGURE 2. Maximum-likelihood phylogram (–InL 73957.608688) of the species of the genus Cnemaspis with Bayesian posterior probabilities (BPP) and maximum-likelihood (ML) bootstrap values, respectively based on the concatenated 3 gene dataset. Black circles are nodes supported by BBP and ML values greater than 0.95 and 70, respectively. Gray circles are nodes supported only by ML values greater than 70. White circles are nodes supported only by BPP values greater than 0.95. The distribution maps on the right delimit the ranges of the four major clades.

Only the following minor discrepancies in the shallow regions of the trees were uncovered between the three loci (see Fig. 2). Both nuclear genes supported the placement of Cnemaspis neangthyi Grismer, Grismer & Thou within a clade of Vietnamese species from the Mekong Delta 364 km to the east and MXRA5 further places one of the specimens within C. siamensis (Smith) from northern Thailand ca. 720 km to the west. However, ND2 supports the placement of C. neangthyi as the sister species to the nearby (25 km) C. chanthaburiensis Bauer & Das. Both nuclear genes also support the placement of C. chanardi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya from the central Thai-Malay Peninsula within this Vietnamese clade whereas ND2 strongly supports its relationship within a clade of other species from the same circumscribed region of southern Thailand. RAG1 indicates that C. huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya from western Thailand is paraphyletic with respect to C. siamensis (Smith) although MXRA5 and ND2 support its monophyly. MXRA5 places an individual of C. peninsularis sp. nov. from Pulau Tioman of the Seribuat Archipelago, Peninsular Malaysia as the sister species to C. shahruli Grismer, Chan, Quah, Muin, Savage, Grismer, Norhayati, Greer & Remegio from northwestern Peninsular Malaysia whereas RAG1 and ND2 place it with other C. peninsularis sp. nov. The nuclear genes were unable to resolve any relationships among the more shallow nodes within the affinis group although nearly all these nodes were supported by ND2 with the topology matching that of the concatenated tree (Fig. 2).

16 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 52 FIGURE 3. Distribution of the species and species groups in the Ca Mau and Northern Sunda clades. Stars indicate type localities.

In deeper regions of the tree, all genes supported the Ca Mau clade as basal to the remaining species of Cnemaspis. RAG1 supported Cnemaspis limi Das & Grismer as the sister species to the nigridia group from northwestern Borneo (Fig. 2) whereas ND2 supports it as the sister species to the kendallii group from southern

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 17 53 Peninsular Malaysia, Borneo, and Indonesia (Fig. 2). MXRA5 remained unresolved with respect to the placement of C. limi. All genes strongly supported the monophyly of the clades and species groups as designated herein as well as the relationships among the four species groups within the Northern Sunda clade (Fig. 2). Because only minor discordances between the three loci were largely restricted to the placement of individual specimens in the more shallow regions of the trees, the data for all loci were concatenated to produce a single, multilocus tree (Fig 2). This tree mirrors the well-supported relationships for the basal nodes as indicated by the nuclear genes and the more shallow relationships revealed by ND2 and essentially bears the same topology as the ND2 tree. Even though the shallow relationships within the affinis group are not well-supported in the concatenated tree (Fig. 2), the topology is supported in the ND2 phylogeny.

FIGURE 4. Distribution of the species in the Pattani and Southern Sunda clades. Stars indicate type localities.

The phylogenetic analyses indicate Cnemaspis is composed of two deeply divergent lineages: one containing the morphologically and behaviorally distinct southern Vietnamese insular endemics C. boulengerii Strauch and C. psychedelica Grismer, Ngo & Grismer referred to herein as the Ca Mau clade and the remaining species of Cnemaspis (Fig. 2). The remaining species form a well-supported lineage containing three major clades referred to as the Pattani, Northern Sunda, and Southern Sunda clades (Fig. 2) whose fragmented distribution along the northern, western and southern fringes of the Sunda Shelf extend from southern Vietnam, Cambodia and Thailand southward through the Thai-Malay Peninsula to Borneo (Figs. 2–4). The Pattani clade is basal to the Northern Sunda and Southern Sunda sister clades (Fig. 2) and restricted to the southernmost portion of peninsular Thailand. The Northern Sunda clade extends from Vietnam to central Peninsular Malaysia and composes a polytomy containing four well-supported, allopatric, species groups referred to as the chanthaburiensis, siamensis, argus, and affinis groups, the latter two being sister lineages (Figs. 2–4). The Southern Sunda clade is conservatively considered a polytomy composed of C. limi of the Seribuat Archipelago; the nigridia group with two species from northwestern Borneo; and the kendallii group containing seven species that extend from southern Peninsular Malaysia and Singapore, eastward through the Seribuat, Anambas, and Natuna Archipelagos to northern Borneo (Figs. 2,4).

18 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 54 FIGURE 5. Ecological parameters of activity period, elevation (upland species are > 600 m in elevation), substrate preference (microhabitat), and the presence or absence of ocelli (eyespots) mapped onto the molecular phylogeny (Fig. 2) of the species of Cnemaspis. Nodes lacking significant statistical support in Figure 2 are collapsed here. Morphological and color pattern synapomorphies are indicated at the appropriate nodes and considered as additional support of the molecular phylogeny. These character states were polarized following the methodology of Maddison et al. (1984) and were not simply mapped onto the tree. Branch lengths do not represent genetic distance.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 19 55 C. mcguirei mcguirei C.

.223 .223 .139 .139 .268 .268 .131 .131 .216 .216 .147 .147 .165 .165 .222 .222 .325 .325 .149 .149 .235 .235 .220 .220 .233 .233 .191 .191 .235 .235 .242 .242 .204 .204 .206 .206 .265 .265 .128 .128 .210 .210 .231 .231

0.000 0.000 C. limi limi C.

0.003 0.003 C. kumpoli kumpoli C.

0.000 0.000 C. kendallii kendallii C. ……continued on the next page page next on the ……continued

0.008 0.008 C. karsticola karsticola C.

0.005 0.005 C. huaseesom huaseesom C.

0.003 0.003 C. harimau harimau C.

0.000 0.000

nov. sp. hangus C.

0.000 0.000 C. grismeri grismeri C.

0.003 0.003 C. flavolineata flavolineata C.

. 2011). Numbers in bold are within Numbers are bold in . 2011). al et divergencespeceis non-bolded and

– C. flavigaster flavigaster C.

0.003 0.003 C. chanthaburiensis chanthaburiensis C.

– C. chanardi chanardi C.

– C. caudanivea caudanivea C.

0.003 0.003 C. boulengerii boulengerii C.

0.005 0.005 C. biocellata biocellata C.

0.055 0.055 C. bidongensis bidongensis C.

0.000 0.000 C. bayuensis bayuensis C.

0.014 0.014 for the mithocondrial gene ND2 calculated in MEGAv5.2.2 (Tamura C. baueri baueri C. 0.000 0.000

Cnemaspis C. aurantiacopes aurantiacopes C.

0.007 0.007 C. argus argus C.

0.016 0.016 C. affinis affinis C.

0.218 0.218 0.267 0.168 0.300 0.268 0.238 0.230 0.352 0.147 0.259 0.164 0.265 0.293 0.204 0.254 0.213 0.247 0.290 0.264 0.233 0.260 0 0.170 0.170 0.133 0.265 0.263 0.146 0.211 0.211 0.345 0.263 0.186 0.278 0.218 0.180 0.124 0.139 0.170 0.241 0.193 0.227 0.201 0.228 0 0.263 0.263 0.223 0.285 0.216 0.273 0.186 0.256 0.340 0.258 0.232 0.253 0.238 0.263 0.253 0.242 0.247 0.254 0.214 0.232 0.253 0.213 0 0.151 0.151 0.186 0.275 0.284 0.022 0.216 0.215 0.332 0.242 0.198 0.273 0.238 0.192 0.117 0.084 0.162 0.225 0.219 0.227 0.210 0.235 0 0.242 0.242 0.231 0.277 0.273 0.232 0.242 0.246 0.353 0.280 0.211 0.289 0.259 0.284 0.222 0.253 0.242 0.189 0.255 0.237 0.227 0.264 0 0.153 0.153 0.205 0.279 0.256 0.185 0.235 0.206 0.342 0.242 0.206 0.304 0.253 0.246 0.127 0.195 0.158 0.260 0.225 0.241 0.224 0.253 0 0.170 0.170 0.215 0.265 0.294 0.115 0.258 0.229 0.345 0.247 0.196 0.273 0.241 0.191 0.146 0.036 0.170 0.249 0.224 0.253 0.247 0.218 0 0.206 0.206 0.229 0.232 0.227 0.211 0.186 0.222 0.379 0.244 0.098 0.273 0.263 0.258 0.191 0.227 0.206 0.235 0.271 0.238 0.227 0.249 0 0.314 0.314 0.361 0.354 0.361 0.344 0.320 0.366 0.198 0.351 0.335 0.366 0.329 0.345 0.302 0.320 0.320 0.354 0.327 0.362 0.392 0.323 0 0.222 0.222 0.178 0.270 0.263 0.191 0.227 0.220 0.345 0.273 0.201 0.294 0.244 0.222 0.137 0.155 0.232 0.216 0.198 0.211 0.206 0.223 0 0.214 0.214 0.116 0.263 0.276 0.231 0.235 0.265 0.335 0.270 0.235 0.284 0.189 0.235 0.219 0.209 0.219 0.253 0.052 0.246 0.250 0.241 0 0.227 0.227 0.207 0.226 0.173 0.209 0.081 0.228 0.335 0.235 0.187 0.236 0.250 0.239 0.209 0.217 0.224 0.242 0.229 0.173 0.186 0.196 0 0.202 0.202 0.210 0.186 0.156 0.235 0.104 0.202 0.334 0.202 0.189 0.220 0.218 0.241 0.213 0.218 0.218 0.206 0.236 0.177 0.202 0.198 0 0.216 0.216 0.177 0.244 0.268 0.187 0.206 0.210 0.340 0.237 0.222 0.289 0.249 0.242 0.191 0.186 0.206 0.235 0.214 0.191 0.170 0.202 0 0.232 0.232 0.244 0.233 0.240 0.235 0.206 0.228 0.379 0.252 0.093 0.299 0.263 0.255 0.209 0.235 0.237 0.235 0.278 0.237 0.216 0.264 0 0.216 0.216 0.253 0.175 0.268 0.266 0.206 0.237 0.353 0.204 0.242 0.186 0.269 0.273 0.240 0.258 0.227 0.277 0.240 0.261 0.237 0.287 0 0.209 0.209 0.206 0.225 0.260 0.205 0.196 0.133 0.361 0.244 0.201 0.232 0.246 0.258 0.198 0.193 0.209 0.182 0.245 0.229 0.131 0.218 0 0.225 0.225 0.186 0.227 0.271 0.218 0.216 0.196 0.329 0.249 0.216 0.259 0.259 0.261 0.201 0.194 0.225 0.220 0.226 0.220 0.198 0.165 0 0.261 0.261 0.247 0.185 0.292 0.270 0.209 0.265 0.344 0.202 0.258 0.165 0.251 0.272 0.263 0.245 0.260 0.273 0.243 0.252 0.243 0.264 0 0.164 0.164 0.197 0.279 0.272 0.146 0.241 0.224 0.327 0.282 0.189 0.303 0.254 0.226 0.109 0.175 0.164 0.208 0.236 0.215 0.226 0.273 0 0.208 0.208 0.216 0.232 0.149 0.212 0.057 0.214 0.336 0.216 0.182 0.247 0.256 0.242 0.208 0.222 0.206 0.244 0.229 0.161 0.198 0.205 0 0.221 0.221 0.241 0.271 0.248 0.239 0.246 0.180 0.352 0.264 0.218 0.277 0.281 0.285 0.215 0.228 0.221 0.238 0.256 0.257 0.212 0.255 0 0.165 0.165 0.186 0.265 0.284 0.146 0.216 0.215 0.314 0.247 0.196 0.289 0.249 0.201 0.039 0.155 0.165 0.225 0.235 0.216 0.216 0.220 0.228 0.228 0.228 0.258 0.236 0.238 0.195 0.230 0.338 0.254 0.226 0.264 0.224 0.246 0.220 0.218 0.218 0.226 0.259 0.218 0.223 0.242 0.242 0.214 0.244 0.247 0.215 0.186 0.158 0.374 0.265 0.232 0.247 0.254 0.278 0.214 0.222 0.232 0.179 0.255 0.206 0.246 0.246 0.234 0.244 0.256 0.235 0.210 0.203 0.362 0.268 0.204 0.239 0.242 0.285 0.213 0.234 0.251 0.191 0.191 0.195 0.259 0.294 0.096 0.237 0.222 0.345 0.253 0.201 0.263 0.226 0.165 0.147 0.041 0.041 0.208 0.270 0.250 0.258 0.123 0.227 0.177 0.250 0.226 0.222 0.255 0.228 0.211 0.236 0.223 0.343 0.229 0.241 0.206 0.276 0.179 0.227 0.343 0.216 0.258 0.262 0.359 0.238 0.229 0.242 0.258 0.265 0.227 0.157 0.207 0.289 0.191 0.240 0.274 0.214 0.273 0.214 0.216 0.245 0.237 0.253 0.211 0.230 0.251 0.000 0.000 0.213 0.265 0.268 0.253 0.251 0.167 0.271 0.201 0.285 0.280 0.206 0.206 0.206 0.216 0.160 0.230 0.258 0.258 0.268 0.238 0.211 0.182 0.258 0.289 0.258 0.205 0.157 0.278 0.268 0.284 0.186 0.232 0.248 0.284 0.259 0.266 0.247 0.194 0.263 0.384 0.283 0.263 0.133 0.187 0.354 0.296 0.216 0.268 0.258 0.376 0.220 0.232 0.314 0.316 0.264 0.222 0.234 0.294 0.191 0.228 0.286 0.228 0.189 0.211 0.211 0.237 0.338 0.223 0.345 0.211 0.280 0.357 0.268 0.216 0.231 0.374 0.170 0.205 0.204 0.344 0.253 0.201 0.348 0.259 0.222 0.381 0.216 0.216 0.244 0.191 0.343 0.220 0.348 0.249 p-distances for the genus

sp. nov. sp. sp. nov. sp. sp. nov. sp. sp. nov. sp. sp. nov. sp. sp. nov. sp. sp. nov. sp.

C. tucdupensis C. tucdupensis C. temiah temiah C. C. sundainsula sundainsula C. C. stongensis C. stongensis C. siamensis siamensis C. C. shahruli C. shahruli C. selamatkanmerapoh C. selamatkanmerapoh C. roticanai roticanai C. C. psychedelica psychedelica C. C. pseudomcguirei pseudomcguirei C. C. perhentianensis C. perhentianensis C. peninsularis C. pemanggilensis pemanggilensis C. C. paripari paripari C. C. omari C. omari C. nuicamensis nuicamensis C. C. niyomwanae niyomwanae C. C. nigridia nigridia C. C. neangthyi neangthyi C. C. narathiawatensis C. narathiawatensis C. mumpuniae C. mumpuniae C. monachorum monachorum C. C. mcguirei mcguirei C. C. limi limi C. C. kumpoli kumpoli C. numbers are between species divergence. C. huaseesom C. huaseesom C. hangus hangus C. C. harimau karsticola C. kendallii C. C. aurantiacopes C. aurantiacopes C. baueri TABLE 4. Uncorrected affinis C. C. argus C. bayuensis C. bidongensis C. bidongensis C. chanthaburiensis chanthaburiensis C. flavigaster C. flavolineata C. grismeri C. C. chanardi C. chanardi C. biocellata biocellata C. boulengerii C. caudanivea C.

20 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 56 C. tucdupensis tucdupensis C.

0.003 0.003

nov. sp. temiah C.

–

nov. sp. sundainsula C.

0.000 0.000

nov. sp. stongensis C.

0.003 0.003 C. siamensis siamensis C.

0.000 0.000 C. shahruli shahruli C.

0.024 0.024 C. selamatkanmerapoh selamatkanmerapoh C.

0.000 0.000 C. roticanai roticanai C.

0.000 0.000 C. psychedelica psychedelica C.

0.000 0.000 C. pseudomcguirei pseudomcguirei C.

0.000 0.000 C. perhentianensis perhentianensis C.

0.012 0.012

. . nov sp. peninsularis C.

0.049 0.049 C. pemanggilensis pemanggilensis C.

0.006 0.006 C. paripari paripari C.

0.000 0.000

nov. sp. omari C.

0.031 0.031 C. nuicamensis nuicamensis C.

0.000 0.000 C. niyomwanae niyomwanae C.

0.010 0.010 C. nigridia nigridia C.

0.014 0.014 C. neangthyi neangthyi C.

0.095 0.095 C. narathiawatensis narathiawatensis C.

0.003 0.003

nov. sp. mumpuniae C.

0.034 0.034 C. monachorum monachorum C. 0.271 0.271 0.255 0.258 0.187 0.269 0.231 0.179 0.256 0.246 0.229 0.244 0.293 0.259 0.347 0.233 0.249 0.241 0.269 0.251 0.267 0.238 0.236 0.236 0.180 0.207 0.262 0.211 0.235 0.148 0.220 0.248 0.255 0.186 0.289 0.188 0.260 0.237 0.206 0.193 0.174 0.180 0.199 0.205 0.229 0.202 0.211 0.193 0.335 0.129 0.247 0.335 0.253 0.186 0.258 0.160 0.278 0.160 0.263 0.237 0.131 0.206 0.224 0.224 0.198 0.141 0.258 0.203 0.190 0.253 0.216 0.172 0.218 0.198 0.214 0.177 0.332 0.192 0.100 0.174 0.234 0.195 0.195 0.231 0.262 0.164 0.249 0.295 0.228 0.238 0.288 0.200 0.294 0.258 0.227 0.207 0.320 0.209 0.237 0.238 0.258 0.243 0.246 0.216 0.227 0.187 0.271 0.328 0.273 0.222 0.356 0.190 0.247 0.268 0.225 0.002 0.002 0.240 0.232 0.266 0.211 0.274 0.218 0.242 0.180 0.259 0.213 0.198 0.268 0.229 0.206 0.238 0.242 0.209 0.170 0.330 0.211 0.230 0.230 0.233 0.220 0.140 0.227 0.201 0.261 0.213 0.246 0.206 0.261 0.261 0.236 0.197 0.231 0.225 0.360 0.248 0.164 0.328 0.218 0.221 0.271 0.320 0.187 0.245 0.191 0.343 0.205 0.237 0.193 0.344 0.256 0.278 0.268 0.369 0.242 0.209 0.214 0.325 0.206 0.236 0.175 0.348 0.237 0.233 0.223 0.381 0.072 0.218 0.316 0.232 0.193 0.352 0.189 0.340 0.183 0.351 0.265 0.206 0.351 0.212 0.212 0.186 0.215 0.253 0.149 0.198 0.258 0.222 0.264 0.264 0.216 0.228 0.256 0.206 0.192 0.223 0.234 0.258 0.275 0.116 0.240 0.249 0.240 0.216 0.208 0.271 0.200 0.218 0.205 0.207 0.197 0.230 0.230 0.075 0.236 0.206 0.215 0.198 0.219 0.206 0.194 0.120

sp. nov. sp. sp. nov. sp. sp. nov. sp. sp. nov. sp.

(Continued) sp. nov. sp. sp. nov. sp. C. tucdupensis C. tucdupensis C. temiah temiah C. C. sundainsula sundainsula C. C. shahruli C. shahruli siamensis C. C. stongensis C. neangthyi neangthyi C. C. monachorum monachorum C. C. mumpuniae C. narathiawatensis C. selamatkanmerapoh TABLE 4. C. nigridia nigridia C. niyomwanae C. C. perhentianensis C. perhentianensis pseudomcguirei C. psychedelica C. roticanai C. C. omari C. omari paripari C. C. nuicamensis nuicamensis C. pemanggilensis C. peninsularis C. Ó

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 21 57 Taxonomy

The molecular analysis corroborates the latest morphological taxonomy proposed for Cnemaspis (Grismer et al. 2010a, 2013a; 2014; Wood et al. 2013) in that no paraphyletic or polyphyletic species had been described. The molecular analysis also supports the recognition of eight new species and a polyphyletic C. kendallii. These issues are addressed in the following species accounts. Additionally, all species groups are provided with morphological diagnoses.

Ca Mau clade

The Ca Mau clade composes the southern Vietnamese insular endemics Cnemaspis boulengerii and C. psychedelica from the Con Dao Archipelago and Hon Khoai Island, respectively, (Figs. 2, 3). Their remarkable color patterns aside, these species are unique among all other Cnemaspis in having caudal tubercles restricted to single paravertebral rows, no lateral caudal furrows, and plate-like femoral and subtibial scales. These character states are considered derived being that they do not occur in any other species of Cnemaspis nor any of the closely related outgroups. Additionally, Grismer et al. (2010b) reported that their behavior is quite atypical from that of other Cnemaspis in that they bask in sunlight and appear to be communal during evening hours. This behavior and their morphology prompted Grismer et al. (2010b) to consider these species closely related which is confirmed here by the molecular and morphological analyses (Figs. 2,5). The genetic divergence between the Ca Mau clade and the remaining species of Cnemaspis (37.6–39.2%; Table 4) is, in many cases, greater than that observed between other gekkonid genera (Gamble et al. 2012). However, we elect not erect a new genus for the 4821 remaining species of Cnemaspis (C. boulengerii Strauch being the type species) in order to maintain taxonomic stability as well as emphasize the monophyletic nature of these two major Southeast Asian lineages. This clade is diagnosed as having a maximum SVL length of 69.0–75.3 mm; 7–10 supralabials; 5–8 infralabials; smooth ventral scales; no precloacal pores; 32–48 paravertebral tubercles; dorsal tubercles linearly arranged and present on flanks; caudal tubercles restricted to a single paravertebral row on each side; no lateral caudal furrows; subcaudals smooth, bearing an enlarged median row; one or two postcloacal tubercles on each side of tail base; enlarged femoral, subtibial and first metatarsal scales; subtibials smooth; 24–32 subdigital lamellae on fourth toe; and dorsal pattern on body, limbs, and tail lacking paired markings or bands.

FIGURE 6. Adult male Cnemaspis boulengerii (LSUDPC 8199) from Con Son Island in the Con Dao Archipelago, Ba-Ria- Vung Tau Province, Vietnam in the dark color pattern phase. Photograph by LLG.

22 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 58 FIGURE 7. Cnemaspis boulengerii from Con Son Island in the Con Dao Archipelago, Ba-Ria-Vung Tau Province, Vietnam. Upper left: adult female (LSUDPC 8219) in the dark color pattern phase. Upper right: adult female (LSUDPC 8200) in the light color pattern phase. Lower left: granite boulder microhabitat on Con Son Island. Lower right: juvenile (LSUDPC 8222) showing the more vivid, light-colored, paravertebral blotches. Photographs by LLG.

Cnemaspis boulengerii Strauch, 1887 Boulenger’s Rock Gecko Figs. 6,7

Gonatodes glaucus Smith 1920:95 (fide Smith 1935:76) Cnemaspis boulengeri Smith 1935:76; Dring 1979:220; Darevsky 1990:128, 1999:34; Grismer et al., 2010b:46

Holotype. Unknown. Type locality: Pulo Condore Island in the South China Sea now Con Dao Island, Ba Ria- Vung Province, Vietnam (fide Sang et al. 2009). Diagnosis. Maximum SVL 69.0 mm; 8–10 supralabials; six or seven infralabials; smooth ventral scales; no precloacal pores; 32–38 paravertebral tubercles; tubercles linearly arranged especially on upper flanks; lateral caudal furrows absent, dorsal caudal furrow weak; caudal tubercles restricted to a single paravertebral row; subcaudals smooth, bearing a medial row of enlarged scales; one postcloacal tubercle on each side; smooth, enlarged, plate-like femoral and subtibial scales; enlarged submetatarsal scales on first toe; 25–32 subdigital fourth toe lamellae; dorsal surfaces unicolor tan; large, subcircular black spots on shoulders and nape; and thin, yellow reticulation on side of neck (Tables 6,7). Color pattern (Figs. 6,7). Dorsal ground color brown to yellowish; dorsum unicolor except for large, black spots on nape, side of neck, and shoulders and a series of elongate, light colored, diffuse, vertebral markings extending onto caudal region and occasionally forming a weak, vertebral stripe; thin, yellowish reticulum on lateral

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 23 59 surfaces of neck and in shoulder regions; ventral surfaces beige, immaculate. Light-colored, vertebral markings in juveniles more prominent, often bearing the typical butterfly shape seen in other species of Cnemaspis. At night the ground color lightens considerably and becomes a dull-yellow. Distribution. Cnemaspis boulengerii is known only from Con Son and Hon Bay Canh islands, Ba Ria-Vung Tau Province, Vietnam of the Con Dao Archipelago, 185 km off the east coast of southern Vietnam (Darevsky 1990, 1999; Grismer et al. 2010b; Fig. 3). It is expected that C. boulengerii occurs on other islands of the archipelago. Natural history. The Con Dao Archipelago contains 16 islands with Con Son Island being the largest and centrally located. Con Son is elongate, hilly, and reaches nearly 570 m in elevation. Much of the low-lying areas of the island are covered in disturbed forest but the rocky, higher elevations of the interior remain fairly intact (Fig. 7) and it is here we observed several specimens of Cnemaspis boulengerii. During the day, C. boulengerii is extremely abundant in lowland forest habitats and can be found climbing on both granite boulders and tree trunks in all planes of orientation. Many lizards were observed basking in dappled light on the tops of boulders and on tree trunks they were seen facing both head up and head down. Lizards were commonly observed in pairs or trios, only found in areas with boulders, and no lizards were observed on the ground. We believe the abundance of this species and its non-secretive nature (unlike that of nearly every other species of Cnemaspis we have observed) may be due to the fact that there are no other diurnal lizards with which to compete. The only other diurnal species observed in their habitat were the skinks Eutropis multifasciata (Kuhl) and Scincella rufocaudata (Darevsky & Nguyen). Cyrtodactylus condorensis (Smith), however, is common at night in the same microhabiats occupied by C. boulengerii during the day. At night, Cnemaspis boulengerii was not seen on the tops of rocks or on the open faces of boulders but was observed only in rock cracks, on the undersides of rocky overhangs, and within caves. Here too, individuals are abundant and commonly found in pairs or trios. Rarely are lizards seen on trees at night and when they are, they are much more wary. Several gravid females carrying two eggs and tens of incubating eggs in communal laying sites in rock cracks and caves were found during August, indicating that this is the reproductive season. Cnemaspis boulengerii is very similar in all aspects of its behavior to its closest relative C. psychedelica (Grismer et al. 2010b). Relationships. Cnemaspis boulengerii is the sister species of C. psychedelica (Fig. 2). Material examined. Vietnam: Ria-Vung Tau Province, Con Dao Archipelago, Con Dao Island CAS 73745, LSUHC 9542, 9578–79, 11364, 11366–68, 11370, MCZ 39014-23, Pulo Condore (= Con Dao), Con Son Island, Vietnam.

Cnemaspis psychedelica Grismer, Ngo & Grismer, 2010 Psychedelic Rock Gecko Fig. 8

Cnemaspis psychadelica Bauer 2013:42.

Holotype. UNS 0444. Type locality: “Hon Khoai Island, Ca Mau Province, Ngoc Hien District, Vietnam (08°26.098 N, 104°49.536 E)” at 30 m in elevation. Diagnosis. Maximum SVL 75.3 mm; 7–10 supralabials; 5–8 infralabials; smooth ventral scales; no precloacal pores; 34–48 paravertebral tubercles; tubercles linearly arranged especially on flanks; lateral caudal furrows absent; caudal tubercles restricted to a single paravertebral row; subcaudals smooth, bearing a medial row of enlarged scales; one or two postcloacal tubercles on each side; smooth, enlarged, plate-like femoral and subtibial scales; enlarged submetatarsal scales on first toe; 24–28 subdigital fourth toe lamellae; occiput and nape bearing a dense, yellow reticulum; hands, feet, forelegs, forelimbs, lower flanks, and tail orange; transverse yellow bars on flanks; ground color of body, brachia, and thighs magenta (Tables 6,7). Color pattern (Fig. 8). Dorsal ground color of head anterior to posterior margin of eyes greenish yellow; occipital region and nape bear a dense, bright-yellow reticulum overlaying thick, black streaks, some of which begin as thin, postorbital stripes; dorsal ground color of trunk and upper limbs immediately proximal to elbow and knee joints blue-gray to magenta; hands, feet, and distal portion of limbs bright-orange; ventral portion of flanks

24 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 60 bright-orange bearing short, transverse, yellow bars; tail bright-orange; all ventral surfaces beige, generally immaculate except for faint stippling on throat and gular region. In alcohol, coloration is generally a uniform dark gray dorsally and beige ventrally, except for a slightly darker gular region.

FIGURE 8. Cnemaspis psychedelica from Hon Khoai Island, Ca Mau Province, Vietnam. Left: adult male (LSUDPC 5062) in the light color pattern phase. Upper right: adult male (LSUDPC 5070) in the dark color pattern phase. Lower right: granite boulder microhabitat on Hon Khoai Island. Photographs by LLG.

Distribution. Cnemaspis psychedelica is known only from Hon Khoai Island, Ngoc Hien District, Ca Mau Province, Vietnam in Rach Gia Bay 18 km off the southern tip of Point Can Mau (Grismer et al. 2010b; Fig. 3). Natural history. Hon Khoai Island is a small (~8 km2) island reaching approximately 320 m in elevation. It slopes moderately to the sea and lacks some of the precipitous, rocky bluffs characteristic of many of the nearby islands. Hon Khoai maintains a thick vegetative cover and is dominated by primary, semideciduous forest on its slopes and upper elevations with disjunct, mangrove swamps fringing its coastlines. Hon Khoai Island’s granite basement gives rise to scattered, small to massive, boulder outcroppings across its lower elevations that provide the microhabitat for Cnemaspis psychedelica (Fig. 8). The vegetation surrounding the granite outcroppings is usually dense and composed of relatively small trees (Grismer et al. 2010b, 2011a). Grismer et al. (2010b) noted that Cnemaspis psychedelica is a relatively large, robust, diurnal, lowland, saxicolous species. Lizards of both sexes and all size classes have been observed abroad on large, granite boulders in the shade of the forest canopy from 0800–1930 hrs. Some lizards were observed basking in filtered sunlight. Lizards would retreat into rock cracks, beneath ledges, or between rocks when threatened. Grismer et al. (2010b) noted it was common to see 2–5 lizards together on the same rock. Lizards showed no preference for any particular plane of orientation, be it vertical, horizontal, or inverted nor did they restrict their activity to only deeply shaded surfaces as do many other species of Cnemaspis (Chan & Grismer 2008; Das & Grismer 2003; Grismer & Chan 2008, 2009; Grismer & Das 2006; Grismer & Ngo 2007; Grismer et al. 2008a,b; 2009). During the evening hours from 1930 to 2400, lizards are not abundant, likely being displaced by the larger Cyrtodactylus sp. 1. At night, the coloration of this species is even more brilliant with the trunk becoming magenta (Fig. 8).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 25 61 Relationships. Cnemaspis psychedelica is the sister species of C. boulengerii (Fig. 2). Remarks. At the time of this writing (22 December 2013), noted Russian reptile dealers are selling illegally collected individuals of Cnemaspis psychedelica for 3500 euro/pair. Unfortunately, the discovery and description of this unique species on this tiny island may ultimately lead to its extinction owing to the wide-reaching criminal element in Southeast Asia. Owing to recent poaching, this species has been put on the IUCN list of threatened species, which ironically will probably increase their commercial value. Material examined. Vietnam: Ca Mau Province, Ngoc Hien District, Hon Khoai Island LSUHC 9254–55, 9257–58, UNS 0444–49 (type series). Additional material examined subsequent to Grismer et al. (2010b): Vietnam: Ca Mau Province, Ngoc Hien District, Hon Khoai Island LSUHC 9524–53, 11007–12.

FIGURE 9. Adult male Cnemaspis monachorum (LSUDPC 6502) from Pulau Langgun of the Langkawi Archipelago, Kedah, Peninsular Malaysia in the dark color pattern phase. Photograph by LLG.

Pattani clade

The Pattani clade is a strongly supported (1.0/100), geographically circumscribed lineage composed of four species from southern Thailand and northwestern Peninsular Malaysia sandwiched between the biogeographic boundaries of the Isthmus of Kra in the north and the Kangar-Pattani Line in the south and embedded within the distribution of the Southern Indochina clade (Fig. 2). The basal species of this clade, C. monachorum Grismer, Norhayati, Chan, Belabut, Muin, Wood & Grismer, is an insular endemic known only from the Langkawi Archipelago of Malaysia. The sister lineage to C. monachorum comprises C. biocellata Grismer, Chan, Nurolhuda & Sumontha from the borderlands of Thailand and Malaysia and the sister species C. niyomwanae Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya from southern Thailand and C. kumpoli Taylor from southern Thailand and extreme northwestern Peninsular Malaysia. The well-supported (0.99/93) sister species relationship between C. niyomwanae and C. kumpoli is further supported in that these are the only species of Cnemaspis to have the derived character states of red bands on their forelimbs and dark-red, dorsal blotches in males (Fig. 5). Three of the four species; Cnemaspis monachorum, C. biocellata, and C. niyomwanae, are small, diurnal, obligate karst-dwellers whereas the remaining species, C. kumpoli, is a much larger nocturnal species that inhabits granite boulders.

26 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 62 This clade is diagnosed in having a maximum SVL of 35.1–63.0; 6–11 supralabials; 5–9 infralabials; smooth ventral scales; 1–12 pore-bearing, precloacal scales with round pores; randomly arranged dorsal tubercles; 2–35 paravertebral tubercles; caudal tubercles not encircling the tail; smooth subcaudals bearing a medial row of enlarged scales; 1–3 postcloacal tubercles on either side of the tail base; no enlarged femoral, subtibial or submetatarsal scales; and 24–41 subdigital lamellae.

FIGURE 10. Cnemaspis monachorum (LSUDPC 6502) from the Langkawi Archipelago, Kedah, Peninsular Malaysia. Upper and middle left: ventral color pattern of male (upper; LSUDPC 8275) and female (middle; LSUDPC 8274) from Pulau Langgun. Photographs by ESHQ. Upper right: adult male (LSUDPC 4701) from Wat Wanaram, Pulau Langkawi in the dark color pattern phase. Lower left: adult female (LSUDPC 4846) from Wat Wanaram in the dark color pattern phase. Lower right: karst microhabitat on Pulau Langgun. Photographs by LLG.

Cnemaspis monachorum Grismer, Norhayati, Chan, Belabut, Muin, Wood & Grismer, 2009 Monks’ Rock Gecko Figs. 9,10

Holotype. ZRC 2.6774. Type locality: “Wat Wanaram, Pulau Langkawi, Kedah, Peninsular Malaysia (06° 20.275 N, 99°52.507 E)” at 35 m in elevation. Diagnosis. Maximum SVL 35.1 mm; seven or eight supralabials; 5–7 infralabials; ventral scales smooth; three, contiguous, pore-bearing precloacal scales with round pores; 2–20 paravertebral tubercles; body tubercles randomly arranged, absent from flanks and lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a medial row of enlarged scales; one or two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 24–30 subdigital fourth toe lamellae; gular region, throat, and abdomen in males yellow; faint, dark, lineate, mid-gular marking present (Tables 6,7). MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 27 63 Singapore Singapore –

et al. Pulau Tenggol Tenggol Pulau

0.009 0.009 Gunung Ledang Ledang Gunung

0.001 0.001 Gunung Panti Panti Gunung

– Selai Selai

0.002 0.002 Pulau Ibol Ibol Pulau

0.002 0.002 Pulau Babi Besar Besar Babi Pulau

0.001 0.001 Pulau Seribuat Seribuat Pulau

0.000 0.000 Pualu Aceh Aceh Pualu 0.002 0.002

sp. nov. for the mithocondrial gene ND2 calculated in MEGAv5.2.2 (Tamura Pulau Tulai Tulai Pulau

– Sungai Lembing Lembing Sungai

– Pulau Tinggi Tinggi Pulau

Cnemaspis peninsularis

0.038 0.038 Pulau Tioman Tioman Pulau

0.007 0.007 Gunung Berlumut Berlumut Gunung

0.003 0.003 Bukit hangus hangus Bukit 0.041 0.041 0.049 0.047 0.075 0.029 0.042 0.074 0.036 0.050 0.052 0.072 0.055 0.080 0.054 0.080 0.090 0.046 0.072 0.038 0.025 0.058 0.090 0.069 0.039 0.042 0.070 0.036 0.042 0.042 0.049 0.046 0.049 0.079 0.031 0.038 0.061 0.039 0.034 0.044 0.009 0.029 0.036 0.050 0.050 0.045 0.003 0.003 0.011 0.056 0.050 0.064 0.091 0.044 0.048 0.074 0.048 0.051 0.064 0.064 0.067 0.059 0.044 0.044 0.053 0.047 0.049 0.081 0.033 0.040 0.065 0.039 0.035 0.046 0.044 0.044 0.052 0.048 0.051 0.079 0.031 0.045 0.063 0.042 0.040 0.045 0.029 0.045 0.045 0.058 0.048 0.056 0.089 0.037 0.046 0.066 0.043 0.044 0.042 0.037 0.037 -distances for the populations of p . Uncorrected 1). Numbers1). in bold are intrapopulational divergencesnon-bolded and numbersare interpopulational divergences. Gunung Panti Gunung Singapore Pulau Tioman Bukithangus Berlumut Gunung TinggiPulau 0.057 Selai Ledang Gunung TenggolPulau TABLE TABLE 5 Sungai Lembing 0.089 0.099 0.101 0.099 201 Tulai Pulau Pulau Seribuat Babi BesarPulau IbolPulau 0.045 0.043 0.026 0.052 0.095 0.037 0.033 0.067 Pualu AcehPualu

28 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 64 kamolnorranathi* kamolnorranathi* 0 0 4 1 0 0 1 1 0 1 1 1 1 0 w w w

7,8 7,8 8,9 8,9 0,1 0,1 1,2 1,2 6,7 6,7 0,w 37.8 37.8

19–24 19–24 24–28 24–28 vandeventeri* vandeventeri* 0 0 3 1 1 0 0 4 0 0 0 1 1 0 0 0 0 1 1 w

8,9 8,9 7–9 7–9 1–3 1–3 44.7 44.7

25–29 25–29 24–28 24–28 punctatonuchalis* punctatonuchalis* / / 0 0 8 5 0 0 0 0 0 1 1 1 0 0 0 0 1 w 7,8 7,8 ant 1–3 1–3

49.6 49.6

24–27 24–27 29–31 29–31 roticanai roticanai page next on the ……continued 9 0 0 8 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 w

7,8 7,8 8,9 8,9 1,2 1,2 3–6 3–6 47.0 47.0

26–2 25–27 25–27

nov. sp. omari 9

0 0 4 1 1 0 1 0 4 0 0 1 1 0 0 1 0 1 0

7,8 7,8 8,9 8,9 w,1 w,0 41.3 41.3

22–2 25–28 25–28 chanardi chanardi 0 0 1 1 0 1 0 1 0 0 0 1 1 0 0 0 0 1 1 25 25 6–8 6–8 6–8 6–8 40.1 40.1 7–10 7–10 20–30 20–30 25–30 25–30

huaseesom huaseesom 0 0 5 0 0 0 0 1 0 0 1 0 1 0 0 1 0 0

1,2 1,2 w,0 6–9 6–9 5–8 5–8 43.5 43.5

7–10 7–10 group 18–24 18–24 21–31 21–31 siamensis siamensis / / 0 0 1 1 0 0 0 1 0 0 1 1 0 0 0 1 1 12 12 8,9 8,9 1,2 1,2

6–8 6–8 39.7 39.7

19–25 19–25 24–26 24–26 siamensis siamensis tucdupensis tucdupensis / / 0 1 0 0 0 0 0 1 1 1 0 0 0 0 0 w 11 11

0-3 0-3 w,1 7–9 7–9 51.0 51.0

8–10 8–10 16–22 16–22 26–32 26–32 nuicamensis nuicamensis 0 0 0 0 0 0 1 1 1 1 0 0 0 0 1 10 10

0,1 0,1 6–7 6–7 7–9 7–9 2–4 2–4 3–6 3–6 48.2 48.2

0, ant ant 0, 0, ant ant 0, 16-21 16-21 27–33 27–33 caudanivea caudanivea 0 9 0 0 1 0 1 1 1 0 0 0 0

8,9 8,9 7,8 1,2 1,2 0,1 0,1 0,w 0,w 0,w 0,w 0–2 0–2 47.2 47.2

0, ant ant 0, 0, ant ant 0, 23-30 23-30 20–24 20–24 aurantiacopes aurantiacopes / / 0 1 1 0 0 0 0 1 1 1 1 0 0 1 0 0 1 17 17

1,2 1,2 58.4 58.4

9–11 9–11 8–10 8–10 23-31 23-31 27–31 27–31 laoensis* laoensis* / / / 0 0 7 9 1 0 1 0 0 1 0 0 1 0 1 0 0 0 w

22 22 29 29 2,3 2,3

40.9 40.9

neangthyi neangthyi group 0 0 5 0 1 0 0 1 2 1 0 1 1 1 0 1 1 1 0 0 1

54.0 54.0

20–26 20–26 22–25 22–25 10–12 10–12 11–13 11–13 chanthaburiensis chanthaburiensis 0 0 8 0 0 0 1 0 1 1 0 1 1 1 0 0 0,1 0,1 0,1 0,1 1–3 1–3 6–9 6–9 42.2 42.2 7–10 7–10 8–10 8–10 21–25 21–25 22–29 22–29

0, post, w 0, post,

Northern Sunda clade chanthaburiensis kumpoli kumpoli 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 13 13

ant 2,3 2,3 7–9 7–9 6–8 6–8 1–8 1–8 63.0 63.0

34-41 34-41 28–35 28–35 niyomwanae niyomwanae 0 0 5 0 0 0 0 3 0 0 0 0 1 0 0 0 0 0 1

1,2 1,2 0,1 0,1 6–8 6–8 56.8 56.8

8–11 8–11 26–31 26–31 31–34 31–34 biocellata biocellata 0 0 0 0 0 0 1 1 0 0 0 1 0 0 1 0 0 1

ant 5–9 5–9 40.1 40.1 25.0 25.0

6–10 6–10 6–12 6–12 21–27 21–27 29–37 29–37 monachorum monachorum 0 0 0 0 0 0 0 3 0 0 0 1 0 0 1 0 0 1 12 12 ant 7,8 7,8 1,2 1,2

5–7 5–7 35.1 35.1 2–20 2–20

24–30 24–30 Pattani clade Pattani boulengerii boulengerii / / / 1 1 0 0 1 1 1 1 0 0 1 0 0 0 0 0 1 from one another. w =weak; ant = anterior; post = posterior; * = species that are not included in the molecular analysis; / = data unavailable.

18 18 6,7 6,7

69.0 69.0 8–10 8–10 32-38 32-38 25-32 25-32 psychedelica psychedelica / / / 1 1 0 0 1 1 1 0 0 1 0 0 0 0 0 1 19 19 1,2 1,2 5–8 5–8 75.3 75.3 7–10 7–10 34–48 34–48 24–28 24–28 Ca Mau clade Cnemaspis Diagnostic morphological characters separating various species of Ventral scales keeled (1) or smooth (0) smooth (0) Ventral scales keeled (1) or No. of precloacal pores (0) separated or (1) continuous pores Precloacal (0) round or (1) elongate pores Precloacal No.ofparavertebral tubercles Tubercles linearly more random (0) arranged (1) or lower flanks on absent (0) Tubercles present (1) or (0) Lateral or absent caudal furrows present (1) (0) not or (1) furrow lateral in tubercles Caudal (0) (1) or not anteriorly caudal tubercles Ventrolateral (0) Lateral row present or absent caudal tubercle (1) paravertebral single a to restricted tubercles Caudal (0) not each or side row on (1) (0) smooth keeled (1)or Subcaudals Single median row of (0) keeledor smooth subcaudals (1) (0) or not encircle tail (1) Caudal tubercles (0) not row (1)Enlarged or median subcaudal scale in males tubercles No. of postcloacal Enlarged femoral scales absent (0) present (1) or (0) or absent scales present (1) subtibial Shield-like smooth (0) scales keeled (1) or Subtibial (1) toe 1st scales on submetatarsal Enlarged (0) not or No. of 4th toe lamellae toe No.of4th Maximum SVL TABLE 6. Samplesize Supralabials Supralabials Infralabials

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 29

65

narathiwatensis narathiwatensis

grismeri grismeri mcguirei mcguirei

……continued on the next page page next on the ……continued

bayuensis bayuensis

nov. sp. stongensis

selamatkanmerapoh selamatkanmerapoh

nov. sp. hangus

flavolineata flavolineata

nov. sp. temiah

shahruli shahruli

affinis affinis harimau harimau

group pseudomcguirei

affinis affinis

perhentianensis perhentianensis

karsticola karsticola argus argus

group flavigaster 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 7 5 5 9 9 10 7 11 7 2 2 2 10 4 19 10 11 1 0 0 0 0,1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 1 0 0,1 0 1 0 1 0 1 0 1 0 1 / 1 0 1 0 1 / 1 0 1 0 1 0 1 0 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0,1 0 w w 1 1 1 1 0,1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0,1 1 1 1 0 0 / 1 1 / / 0 0 0,1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 1 0 1 0 0 0 1 1 1 0 1 ant 0, 0 0 0 0,ant ant 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0,w 1 0 w 0 0 0 0 0 0 1 0 1 0 1 0 1 0 ant 0 1 0 1 0 1 0 1 0 ant 1 0 1 0 ant 0,1 0 1 0 1 1 1 1 1 7,8 7,8 6–10 7,8 6–8 1–5 4 5,6 0 5–7 5,6 0 1 5–8 5–9 5–10 8–10 3–6 1,2 1,2 1–4 2,3 3–4 2,3 2,3 2 1–3 2,3 2 2 3 2,3 2 2–5 2,3 1–3 50.1 50.1 65.2 48.1 47.0 42.5 40.7 50.8 36.5 46.7 39.2 50.5 43.4 49.3 46.1 65.0 50.6 53.2 9,10 9,10 8,9 7,8 8–10 9,10 9,10 9–13 10,11 8–10 9 9 10 8–11 9,10 7–10 8 9,10 8–10 8–10 8,9 6,7 7–8 8–10 9,10 8–10 8–10 7–10 9 8 9,10 8–10 8,9 7–9 8 7–11 21–24 21–24 26–32 17–19 22–27 23–32 18–20 20–28 19–23 22–27 23 22–24 30 26–33 23–30 26–32 27–32 28–34 29–34 29–34 31–35 27–30 28–31 23–26 25–30 28,29 21–30 22–26 23 27–34 31–33 28–32 27–32 27–35 25–31 24–30 Northern Sunda clade argus (Continued) (Continued) Maximum SVL TABLE 6. TABLE Supralabials Infralabials Ventral scales keeled (1) or smooth (0) smooth (0) Ventral scales keeled (1) or Tubercles linearly more random (0) arranged (1) or lower flanks on absent (0) Tubercles present (1) or No. of precloacal pores No. of precloacal pores No.ofparavertebral tubercles (0) Lateral or absent caudal furrows present (1) Precloacal pores continuous (1) or separated (0) (0) separated or (1) continuous pores Precloacal (0) (1) or round elongate Precloacal pores (0) not or (1) furrow lateral in tubercles Caudal Ventrolateral caudal tubercles anteriorly (1) or not (0) (0) not or (1) anteriorly tubercles caudal Ventrolateral (0) Lateral row present or absent caudal tubercle (1) paravertebral single a to restricted tubercles Caudal (0) not each or side row on (1) (0) smooth keeled (1) or Subcaudals (0) not row (1)Enlarged or median subcaudal scale in males tubercles No. of postcloacal Enlarged femoral scales absent (0) present (1) or (1) toe 1st scales on submetatarsal Enlarged (0) not or lamellae toe No.of4th Samplesize Single median row of keeled subcaudals (1) or smooth (0) Singlemedian(0) row of keeled orsmooth subcaudals (1) (0) not or (1) tail encircle tubercles Caudal (0) or absent scales present (1) subtibial Shield-like smooth (0) scales keeled (1) or Subtibial

30 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 66

* * nov. sp. sundagekko

nov. sp. peninsularis

bidongensis bidongensis

baueri baueri

pemanggilensis pemanggilensis

nov. sp. mumpuniae

nov. sp. sundainsula

group kendallii kendallii

kendallii kendallii

dringi* dringi* paripari paripari

group nigridia nigridia

nigridia nigridia limi limi / 0 0 0 / / / / / / / / / 0 0 0 / / / / / / / / 0 1 1 0 0 w,1 0 0 0 0 0 0 0 10–16 2–6 5,6 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 1 1 1 0 0,w 0 0 w 1 w,0 0 0 0 w,0 0,w 0 1 1 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 w w,1 1 1 1 1 1 1 1 1 1 1 w 1 1 1 1 1 1 1 1 1 1 1 w 1 1 1 0 0,w,post 1 1 1 1 0 post w 0 0 0 w,1 1 w,1 0,w w,0 0 w,1 0,w 34 34 4 5 2 14 18 17 18 27 14 86 6 1,2 1,2 2–4 2 2 2 2–4 1,2 1,2 1,2 1,2 1,2 2,3 88.2 88.2 75.5 50.7 46.4 58.4 84.5 60.9 76.0 67.4 58.1 60.0 68.0 8–12 8–12 10,11 12,13 11 10,11 8–11 9–12 10–13 11–13 9,10 10,11 11–13 7–10 7–10 9–11 10,11 9 8,9 7–10 8–11 8–10 8–12 7–9 7–10 8–11 25–35 25–35 39–43 26–31 25–27 18–26 26–37 18–25 30–37 18–27 21–26 17–25 20–25 29–36 26–29 26–31 32–35 25–33 25–29 29–35 27–34 26–32 26–30 27–33 33–38 Southern Sunda clade (Conintued) (Conintued) Single median row of keeled subcaudals (1) or smooth (0) Single median row of (0) keeledor smooth subcaudals (1) Subcaudals keeled (1) or smooth (0) (0) smooth keeled (1) or Subcaudals (0) not or (1) tail encircle tubercles Caudal smooth (0) scales keeled (1) or Subtibial (1) toe 1st scales on submetatarsal Enlarged (0) not or Lateral caudal furrows present (1) or absent (0) (0) Lateral or absent caudal furrows present (1) (0) not or (1) furrow lateral in tubercles Caudal (0) not or (1) anteriorly tubercles caudal Ventrolateral (0) Lateral rowpresent or absent caudal tubercle (1) paravertebral single a to restricted tubercles Caudal (0) not or each side (1) row on (0) not row (1)Enlarged or median subcaudal scale in males tubercles No. of postcloacal (0) or absent scales present (1) subtibial Shield-like lamellae toe No.of4th Infralabials smooth (0) Ventral scales keeled (1) or No. of precloacal pores (0) separated or (1) continuous pores Precloacal (0) round or (1) elongate pores Precloacal Tubercles linearly more random (0) arranged (1) or lower flanks on absent (0) Tubercles present (1) or Enlarged femoral scales absent (0) present (1) or Samplesize Supralabials Supralabials No.ofparavertebral tubercles Maximum SVL TABLE 6. TABLE

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 31 67 TABLE 7. Diagnostic color pattern characters separating various species of Cnemaspis from one another. Var = charcter variable; / = data unavailable. Ca Mau Pattani clade Northern Sunda clade clade chanthaburiensis group siamensis group argus group

sis sp.nov. argus argus kumpoli chanardi roticanai laoensis* siamensis neangthyi neangthyi karsticola biocellata flavigaster huaseesom huaseesom caudanivea boulengerii tucdupensis nuicamensis niyomwanae psychedelica monachorum vandeventeri* aurantiacopes omari perhentianensis perhentianensis chanthaburien kamolnorranathi* punctatonuchalis*

Dorsal color pattern sexually dimorphic no no no yes yes yes no no / yes no no no no yes no no yes yes no no no no no no Ventral pattern sexually dimorphic no no yes yes / no yes no / yes no no no yes yes yes / yes yes yes / yes no no no Head yellow no no no no no no no no no no no no no no yes no no no no no no no no no no Reddish blotches on head and body no no no no no yes no no no no no no no no no no no no no no no no no no no Dense yellow reticulum on occiput and side of neck yes no no no no no no no no no no no no no no no no no no no no no no no no Ocelli on occiput and nape no no no yes no no no no no no no no no no no no no no no no no no no no no Ocelli on shoulder no no no yes no yes no no no no no no no no no no no no no no no no no no no Dual ocelli on shoulder no no no no no no no no no no no no no no no no no no no no no no no no no Ocelli on brachium and side of neck no no no no no no no no no no no no no no no no no no yes no no no no no no Thin, white, nuchal loop no no no no no no no no no no no no no no no no no no no no no no no no no Large, black round spots on nape and anterior of body no yes no no no no no no no no yes no no no no no no no no no no no no no no Thin yellow reticulum on side of neck no yes no no no no no no no no no no no no no no no no no no no no no no no Yellowish, prescapular crescent no no no no no no no no no no no no no no no yes yes yes no yes var no no no no Forelimbs yellow no no no no no no no no no no no no no no yes no no no no no no no no no no Hind limbs yellow no no no no no no no no no no no no no no yes no no no no no no no no no no Reddish blotches or bands on limbs no no no no yes yes no no no no no no no no no no no no no no no no no no no Forearms and forelegs orange yes no no no no no no no no yes no no no no no no no no no no no no no no no Dorsal ground color magenta yes no no no no no no no no no no no no no no no no no no no no no no no no Dorsal ground color reddish no no no no no no no no no no no no no no no no no no no no no no no no no Uniform brown ground color no yes no no no no no no no no no no no no no no no no no no no no no no no Light vertebral stripe no no no no no no no no no no no no no no no no no no no no no no no no no Yellow postscapular band no no no no no no no no no no no no no no no no no no no no no no no no no Black, squarish, paired, paravertebral dorsal markings no no no no no no no no no no no no yes no no no no no no no no no no no no Small, light, round spots on flanks no no no no no no no no no no no no no no no no no no no no no no no no no Black flanks with distinct yellowish spots no no no no no no no no no no no no no no no no no no no no no no no no no Yellow or white bars on flanks yes no no no no no no no no no no no no no no no no no no no no no no no no Original tail yellow no no no yes no no no no no no no no no no var no no no no no no no no no no Original tail orange yes no no no no no no no no yes no no no no no no no no no no no no no no no Regenerated tail yellow no no no no no no no no / no / no no no no no no yes / no no no no no no Regenerated tail orange yes no no yes no no no no / no / no no no no no no no / no no no no no no White, dorsal caudal tubercles no no no no no no no no no no no no no no no no no no no no no no no no no Caudal bands present no no yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes no no no no Wide black and yellow bands on tail no no no no no no no no no no yes no yes no no no no no no no no no no no no Thin, yellow caudal bands anteriorly no no no no no no no no no no no no no no no no no no no no no no no no no Posterior portion of original tail white no no no no no no no no no no yes no no no no no no no no no no no no no no Posterior portion of original tail black no no no no no no no no no no no no yes no no no no no no no no no no no no Disticnt black and white caudal bands at least posteriorly no no no no no no no no no no no no no no no no no no no no no yes yes no yes Gular region orange no no no no no no yes no / yes no no yes no no no no no / yes no no no no no Gular region yellow no no yes no no / no no / no no no no yes yes yes yes yes / no no no no no no Lineate gular markings no no yes no no no no no / yes no yes no yes no no no no / no no no no no no Throat yellow no no yes no no / no no / no no no no yes yes no no no no no no no no no no Throat orange no no no var no no yes no / yes no no yes no no no no no yes no no no no no no Pectoral region yellow no no yes no no / no no / no no no no yes yes no no yes / no no no no no no Pectoral region orange no no no var no no no no / yes no no yes no no no no no / var no yes no no no Abdomen yellow no no yes var no / no no / no no no no no no yes yes yes / no no no no no no Abdomen orange no no no var no no yes no / yes no no yes no no no no no / yes no yes no no no Ventral surfaces of forelimbs orange no no no no no no no no / yes no no yes no no no no no / yes no no no no no Ventral surfaces of forelimbs yellow no no no no no no no no / no no no no no yes no no no / no no no no no no Ventral surfaces of hind limbs orange no no no var no no no no / yes no no yes no no no no no / yes no yes no no no Ventral surfaces of hind limbs yellow no no no no no no no no / no no no no no no yes yes yes / no no no no no no Subcaudal region yellow no no no var no no no no / no no no no no yes yes yes yes no no no no no no no Subcaudal region orange no no no var no no yes no / no no yes no no no no no no yes yes no yes no no no At least posterior half of subcaudal region white no no no no no no no no no no yes no no no no no no no no no no no no yes no ...... continued on the next page

32 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 68 TABLE 7. (Continued) Northern Sunda clade Southern Sunda clade affinis group nigridia group kendallii group

*

sp. nov. sp.nov. sp. nov. sp.nov. sp. nov. sp.nov. sp.nov. limi affinis affinis baueri baueri dringi* dringi* nigridia shahruli paripari paripari harimau grismeri grismeri kendallii mcguirei bayuensis bidongensis bidongensis flavolineata temiah pemanggilensis hangus hangus pseudomcguirei narathiwatensis stongensis mumpuniae selamatkanmerapoh sundainsula peninsularis peninsularis sundagekko

Dorsal color pattern sexually dimorphic yes yes yes yes no no no no no no yes yes yes no no yes / no no no no no yes yes / Ventral pattern sexually dimorphic no yes no yes no no no no no no no no no no no yes / no yes no no no no yes / Head yellow no no no no no no no no no no no no no no no yes no no no no no no no no no Reddish blotches on head and body no no no no no no no no no no no no no no no no no no no no no no no no no Dense yellow reticulum on occiput and side of neck no no no no no no no no no no no no no no no no no no no no no no no no no Ocelli on occiput and nape no no no no no no no no no no no no no no no no no no no no no no no no no Ocelli on shoulder no yes yes yes no no no no no no yes yes yes no no no no no no no no no no no no Dual ocelli on shoulder yes no no no no no no no no no yes yes no no no no no no no no no no no no no Ocelli on brachium and side of neck no no no no no no no no no no no no no no no no no no no no no no no no no Thin, white, nuchal loop no no no no no no no no no no no no no no no no no no no yes no no no no no Large, black round spots on nape and anterior of body no no no no no yes no no no no no no no yes yes no no no no no no yes yes no yes Thin yellow reticulum on side of neck no no no no no no no no no no no no no no no no no no no no no yes no no no Yellowish, prescapular crescent no no no no no no no no no no no no no no no no no no no no no no no no no Forelimbs yellow no no no no no no no no no no no no no no no yes no no no no no no no no no Hind limbs yellow no no no no no no no no no no no no no no no yes no no no no no no no no no Reddish blotches or bands on limbs no no no no no no no no no no no no no no no no no no no no no no no no no Forearms and forelegs orange no no no no no no no no no no no no no no no no no no no no no no no no no Dorsal ground color magenta no no no no no no no no no no no no no no no no no no no no no no no no no dorsal ground color reddish no no no no no no no no no no no no no no no no no no no yes no no no no no Uniform brown ground color no no no no no no no no no no no no no no no no no no no no no yes no no no Light vertebral stripe var no no var var var no no no no no no var no no no no no no no no no no no no Yellow postscapular band no var var no no no no no no no yes yes no no no no no no no no no no no no no Black, squarish, paired, paravertebral dorsal markings no no no no no no no no no no no no no no no no no no no no no no no no no Small, light, round spots on flanks no no no no no no no no no no no no no no no no yes no yes yes yes no no no no Black flanks with distinct yellowish spots no no no no no no no no no no no no no no no no no no no no no no no no no Yellow or white bars on flanks yes yes yes no no no no no yes yes yes yes yes no yes no no no no no no no no no no Original tail yellow no no no no no no no no no no no no no no no no no no no no no no no no no Original tail orange no no no no no no no no no no no no no no no no no no no no no no no no no Regenerated tail yellow no no no no no no no no no no no no / no no yes no no no yes no no no yes / Regenerated tail orange no no no no no no no no no no no no / no no no no no no no no no no no no White, dorsal caudal tubercles no no no yes no no no no no no no no no yes no no no no no no no yes no no yes Caudal bands present no no no no no no no no no no no no no no no no no no no no no no no no no Wide black and yellow bands on tail no no no no no no no no no no no no no no yes no no no no no no no no no no Thin, yellow caudal bands anteriorly no no no no no no no no no no no no no no no no no no no no no yes no no no Posterior portion of original tail white no no no no no no no no no no no no no no no yes no no no no no no no no no Posterior portion of original tail black no no no no no no no no no no no no no no no no no no no yes no yes no yes no Disticnt black and white caudal bands at least posteriorly no yes no var no no no no yes no yes yes yes no no no / yes no no no no yes, f no yes Gular region orange no no no no no no no no no no no no no no no no / no no no no no no no no Gular region yellow no yes yes yes no no no no no no no no no no no no / no yes no no no no no no Lineate gular markings no no / no no no no no no no no no no no no no / no no no no no no no no Throat yellow no yes / yes no no no no no no no no no no no no / no yes no no no no no no Throat orange no no / no no no no no no no no no no no no no / no no no no no no no no Pectoral region yellow no no / yes no no no no no no no no no no no no / no no no no no no no no Pectoral region orange no no / no no no no no no no no no no no no no / no no no no no no no no Abdomen yellow no no / no no no no no no no no no no no no no / no yes no no no no no no Abdomen orange no no / no no no no no no no no no no no no no / no no no no no no no no Ventral surfaces of forelimbs orange no no / no no no no no no no no no no no no no / no no no no no no no no Ventral surfaces of forelimbs yellow no no / no no no no no no no no no no no no no / no no no no no no no no Ventral surfaces of hind limbs orange no no / no no no no no no no no no no no no no / no no no no no no no no Ventral surfaces of hind limbs yellow no no / no no no no no no no no no no no no no / no no no no no no no no Subcaudal region yellow no no / no no no no no no no no no no no no no / no yes no no no no no no Subcaudal region orange no no / no no no no no no no no no no no no no / no no no no no no no no At least posterior half of subcaudal region white no no no no no no no no no no no no no yes no yes no yes yes no no no no no no

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 33 69 Color pattern in life (Figs. 9,10). Dorsal ground color brown; head, body, and limbs overlain with small, irregularly shaped, somewhat randomly arranged, black and cream colored spots varying somewhat in size; light dorsal blotches in general paravertebral arrangement; markings on top of head smaller than those on body; dark markings in caudal region tend to form bands that encircle tail; anterior gular region yellowish, most pronounced on mental scale; faint, dark, midgular stripe present; pectoral region and anterior, abdominal region orangish yellow; remainder of ventral surfaces of body and limbs beige bearing faint, dark, stippling. Distribution. Cnemaspis monachorum is known only from the Langkawi Archipelago where it inhabits the islands of Langkawi (Grismer et al. 2009) and reported here for the first time from the smaller satellite island of Langgun (Fig. 4). Natural history. On Pulau Langkawi, Grismer (2011a) noted that Cnemaspis monachorum is a lowland, saxicolous species known only from the karst outcropping of Wat Wanaram near the town of Kuah in a region dominated by a mixture of primary coastal and lowland dipterocarp forest. Cnemaspis monachorum is a small, swift, agile species abroad only during the day, climbing on the fragmented boulders along the periphery of karst formations as well as along the base of karst cliffsides near the edges of cracks. Lizards are wary and will rapidly retreat into a crack while curling their tail above their back and waving it from side to side. During the night, lizards are not abroad and only occasionally observed deep within limestone cracks. Cnemaspis monachorum is the smallest species of Cnemaspis and possibly associated with this small size is that gravid females carry only a single egg whereas females of all other Cnemaspis carry two eggs. Gravid females have been observed in October. On Pulau Langgun, C. monachorum is also diurnal and common throughout the island on the limestone rocky hillsides as well as the cliff faces that edge Tasik (lake) Langgun (Fig. 10). During August, no gravid females were found on Pulau Langgun suggesting that October may be the beginning of the reproductive season. Relationships. Cnemaspis monachorum is the basal lineage of the Pattani clade (Fig. 2). Material examined. Malaysia: Kedah; Pulau Langkawi LSUHC 9118, ZRC 2.6774–76 (type series). Material examined since Grismer et al. (2009): Malaysia: Kedah; Pulau Langkawi LSUHC 9115–17; Pulau Langgun LSUHC 10807–11.

Cnemaspis biocellata Grismer, Chan, Nurolhuda & Sumontha, 2008 Twin-spot Rock Gecko Figs. 11,12

Holotype. ZRC 2.66693. Type locality: “Kuala Perlis, Perlis, Peninsular Malaysia (06°24.437N, 100°08.564E) at 37 m in elevation. Diagnosis. Maximum SVL 40.1 mm; 6–10 supralabials; 5–9 infralabials; ventral scales smooth; 6–12 contiguous, pore-bearing precloacal scales with round pores; 21–27 paravertebral tubercles; body tubercles randomly arranged, present on flanks, absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a median row of enlarged scales; one postcloacal tubercle on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 29–37 subdigital fourth toe lamellae; paired ocelli on occiput in males; wide, white to yellow nuchal loop in males; single ocellus in shoulder region in males; original tail yellow and regenerated tail orange in males; and throat, pectoral region, ventral surface of hind limbs, abdomen, and subcaudal region orange or yellow in males (Tables 6,7). Color pattern in life (Figs. 11,12). Adult males: ground color of dorsal surface of head, body, limbs, and tail dull yellow; rostrum grayish with faint, light markings highlighting bright yellow, anterior, extra brillar fringe; interorbital region yellow with two, distinct, white, immaculate, well-defined occipital ocelli; ocelli accentuated by wide, black, occipital band forming anterior border of a series of closely spaced, large, white to yellow spots forming a nuchal band extending from posterior margin of one eye to posterior margin of other eye; small, black, shoulder patch enclosing a single, white to yellow ocellus; body overlain by five yellow, butterfly-shaped, vertebral blotches extending from shoulder region to base of tail; small, faint yellow blotches on flanks and limbs; blotches tend to form caudal bands; ventral surfaces yellow or orange, nearly immaculate with only a faint amount of subcaudal mottling. Adult females: ground color of dorsal surfaces light brown lacking black occipital and shoulder markings of males; occiput bears faint, straw colored, ocelli homologous to those in males; poorly

34 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 70 defined, straw colored, butterfly-shaped, vertebral markings extend from nape to base of tail continuing posteriorly to form poorly defined, caudal bands; small, irregularly shaped, faint spots on flanks and limbs; and Ventral surfaces beige, nearly immaculate with only faint, subcaudal mottling.

FIGURE 11. Cnemaspis biocellata from Perlis, Peninsular Malaysia. Upper left and right: adult male (LSUDCP 4231) in the dark color pattern phase and adult female (LSUDPC 4230) in the light color pattern phase, respectively, from Gua Kelam. Lower right: adult male (LSUDPC 4233) in the dark color pattern phase from Kuala Perlis. Middle left: adult female (LSUDPC 5949) in the dark color phase from Wang Kelian. Photographs by LLG. Lower left: adult male (LUSDPC 6247) from Perlis State Park in the dark color pattern phase. Photograph by ESHQ.

Distribution. Cnemaspis biocellata extends through the karst system of the Banjaran Nakawan from Thale Ban National Park, Satun Province, Thailand southward through Wang Kelian and Perlis State Park to Gua Kelam, Tasik Meranti, Kampung Bukit Cabang, and Kuala Perlis, Perlis in northern Peninsular Malaysia (Grismer 2011a; Fig. 4). Natural history. Grismer (2011a) noted that Cnemaspis biocellata occurs in lowland areas from near sea level to approximately 200 meters in elevation and appears to be a karst-substrate specialist. Lizards have been observed on trees and cement walls (Fig. 11) but only where they were adjacent to a karst formation. Cnemaspis biocellata is often observed abroad during the day on the shaded, vertical surfaces of karst walls and boulders as well as within crevice microhabitats and beneath small limestone rocks piled on the ground. During the day, lizards are wary and difficult to approach and will retreat into nearby crevice microhabitats at the slightest provocation. At night, lizards move farther away from these retreats and are more approachable. At Gua Kelam, Perlis the habitat is highly disturbed and continually frequented by visitors to the neighborhood park yet C. biocellata are abundant and easily observed both day and night. At Tasik Meranti, Perlis, lizards occur on large karst rocks in lowland dipterocarp forest but restrict their daytime activities to shaded areas. At Wang Kelian, they are abroad only during the day, perhaps due to the presence of the much larger Cyrtodactylus astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer & Pauwels and Gekko gecko Linnaeus at night which may

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 35 71 even prey on them. At Kampung Bukit Cabang, lizards have been found at night within caves. At Kuala Perlis, lizards are abundant during the day and night on karst formations in the vicinity of parks, parking lots, and housing communities in areas with no immediate native vegetation. These observations suggest that the most important environmental component for this species is the karst substrate and the microhabitats it offers, regardless of the condition of the surrounding forest. This small species is amazingly quick and agile and effortlessly moves from one inclined surface to another. Upon capture, lizards release large sections of their skin in much the same manner as the Stump-tailed Gecko, Gehyra mutilata (Wiegmann). Cnemaspis biocellata is the second smallest species of Cnemaspis and females carrying one or two eggs have been observed only during March. Relationships. Cnemaspis biocellata is a member of the Pattani clade and most closely realeted to the sister species C. niyomwanae and C. kumpoli (Fig. 2). Material examined. Malaysia: Perlis, Kuala Perlis ZRC 2.6693–98 (type series). Specimens examined since Grismer et al. (2008a): Malaysia: Perlis, Gua Kelam LSUHC 8787–92, 8802, 8804–05, 9682. Kampung Bukit Chabang LSUHC 9683–84.

FIGURE 12. Cnemaspis biocellata from Perlis, Peninsular Malaysia. Upper right; adult female (LSUDPC 4229) from Gua Kelam in the light color pattern phase. Lower right: variation in ventral coloration in adult males from Bukit Cabang (LSUDPC 5338). Left: karst microhabitat at Wang Kelian. Photographs by LLG.

Cnemaspis niyomwanae Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010a Niyomwan’s Rock Gecko Fig. 13

Holotype. THNHM 15910. Type locality: “Thum Khao Ting, Palean District, Trang Province, Thailand (07°09.943N, 99°48.142E) at 28 m in elevation.”

36 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 72 FIGURE 13. Cnemaspis niyomwanae from Thum Khao Ting, Palean District, Trang Province, Thailand. Upper left: adult female (LSUDPC 7035) in the light color pattern phase. Upper right: adult male (LSUDPC 7032) in the dark color pattern phase. Middle left: ventral coloration of adult female (LSUDPC 7036) which is the same as adult males. Middle right: subadult (LSUDPC 7034) in the light color pattern phase. Lower: habitat at Thum Khao Ting. Photographs by MS.

Diagnosis. Maximum SVL 56.8 mm; 8–11 supralabials; 6–8 infralabials; ventral scales smooth; three usually contiguous, pore-bearing precloacal scales with round pores; 26–31 paravertebral tubercles; body tubercles randomly arranged, absent from flanks and from lateral caudal furrows; no ventrolateral caudal tubercles; no lateral row of caudal tubercles; caudal tubercles not encircling tail; subcaudals smooth bearing a median row of enlarged scales; one or two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 31–34 subdigital fourth toe lamellae; and reddish bands on limbs in males (Tables 6,7).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 37 73 Color pattern in life (Fig. 13). Dorsal ground color of head, body, and tail faded green; dorsal ground color of limbs faded brown; yellow stripe on each canthus rostralis; diffuse, light, paired, occipital blotches present; faint, light mottling on sides of head; whitish, medial blotch on nape followed posteriorly by five, lightly colored, paravertebral, butterfly-shaped markings between forelimb insertions and base of tail; markings continue onto tail to form lightly colored bands; enlarged, white tubercles on sides of neck, shoulders and flanks; other tubercles on body dark or lightly colored; faint, reddish dorsal blotches on body; upper regions of limbs bearing diffuse light mottling; alternating red and yellow bands on forelimbs and forelegs; digits white bearing broad, brown bands; all ventral surfaces except subcaudal region of uniform beige with fine, dark stippling in some scales; and subcaudal region grayish. Distribution. Cnemaspis niyomwanae is known only from the border regions of Trang and Satun Provinces, Thailand (Fig. 4). Natural history. Very little is known about the life history of Cnemaspis niyomwanae. Grismer et al. (2010a) reported that it occurs in lowland karst areas in the vicinity of small streams (Fig. 13) and that it is abroad at night. During the day, lizards retreat into limestone crevices and caves. Relationships. Cnemaspis niyomwanae is a member of the Pattani clade and the sister species of C. kumpoli Taylor (Fig. 2). Material examined. Thailand: Trang Province, Palean District, Thum Khao THNHM 15910. La-ngu District, Baan Man Pud CUMZ R-2009, 6,24-10, KZM 008, PSUZC-RT 2010.56, ZMKU Re-000315. These specimens represent the type series.

Cnemaspis kumpoli Taylor, 1963 Kumpol’s Rock Gecko Fig. 14

Holotype. FMNH 178268. Type locality: “Khao Chong, Forestry Experimental Station, Trang province, Thailand.” Diagnosis. Maximum SVL 63.0 mm; 7–9 supralabials; 6–8 infralabials; ventral scales smooth; 1–8, discontinuous, pore-bearing precloacal scales with round, poorly developed pores; 28–35 paravertebral tubercles; body tubercles randomly arranged, present on flanks; tubercles in lateral caudal furrows anteriorly; no ventrolateral caudal tubercles; no lateral caudal row of tubercles; caudal tubercles not encircling tail; subcaudals smooth bearing a median row of enlarged scales; two or three postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 34–41 subdigital fourth toe lamellae; single ocellus in shoulder region in males; red bands on forelimbs and hind limbs in males; and reddish blotches on dorsum and tail in males (Tables 6,7). Color pattern (Fig. 14). Adult males: dorsal ground color lime-green to yellow, overlain by red blotches on head (usually), body, limbs, and tail; round, red to brownish, paravertebral markings extend from just posterior to forelimb insertions to base of tail alternating with smaller, yellow, paravertebral blotches; smaller, more irregularly shaped, red blotches occur on flanks and limbs, those on tail tend to form bands; poorly defined, black markings occur on anterior margin of nape highlighting a white, nuchal band between it and large, black, shoulder patches; a single, whitish, longitudinal bar enclosed in each shoulder patch; shoulder patches narrowly meet on midline of body; ventral surfaces beige, immaculate; subcaudal region faintly mottled. Adult females and juveniles: ground color dull yellow; no red or brownish markings on head or limbs; no black shoulder patches enclosing ocelli; yellow markings on head anteriorly; paired, symmetrical dark and light markings on occiput; a series of white, vertebral blotches alternating with paired, dark vertebral blotches extend from nape to base of tail then transforming into indistinct, caudal bands; regularly shaped, dark and light markings on flanks and limbs; ventral surfaces beige, immaculate; faint, subcaudal mottling pattern present. The illustration of Cnemaspis kumpoli in Das (2010:57) is misleading and the description of this species’ color pattern (Das 2010:202) does not take into account its marked sexual dimorphism. Distribution. Cnemaspis kumpoli ranges from southern Thailand south of the Isthmus of Kra from the Khao Chong Forest Reserve, Trang, Satun, and Songkhla provinces, southwestern to extreme northwestern Malaysia where it is known only from Kaki Bukit, Perlis and Perlis State Park along the Thai-Malaysian border (Grismer 2011a; Grismer et al. 2010a; Fig. 4).

38 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 74 FIGURE 14. Cnemaspis kumpoli. Upper left and right: adult female (LSUDPC 4222) and adult male (LSUDPC 4220), respectively, from Perlis State Park, Perlis, Peninsular Malaysia in the dark color pattern phase. Lower left: granite boulder microhabitat at Perlis State Park. Photographs by LLG. Lower right: adult male (LSUDPC 8271) from Songkhla, Songkhla Province, Thailand in the light color pattern phase. Photograph by MS.

Natural history. In Peninsular Malaysia, Cnemaspis kumpoli occurs in rocky areas composed of granite boulders within primary, lowland dipterocarp forest and has not been observed on nearby karst formations (Grismer 2011a). Lizards are seen at night on large boulders of granite outcroppings on steep hillsides that often border streams. They are quite active, remain wary, and do not venture far from safe retreats between the rocks, within rock cracks, or from near burrows at the base of the rocks. Grismer (2011a) reported finding a pair of adults on the base of a large tree near granite boulders. No specimens have been observed abroad during the day and females carrying two eggs have been reported during June and September (Grismer 2011a). Relationships. Cnemaspis kumpoli is a member of the Pattani clade and the sister species of C. niyomwanae (Fig. 2). Remarks. Despite the fact that Cnemaspis kumpoli is a large, nocturnal, granite dwelling species embedded in a clade of small, diurnal karst-dwellers it bears none of the characteristics seen in most other large granite dwelling species (i.e., C. argus, C. limi, C. mcguirei and C. perhentianensis) such as keeled ventrals, keeled subtibials, and keeled subcaudal scales and strong dorsal tuberculation. Instead, it is weakly tubreculated and has smooth ventrals, subtibtials, and subcaudal scales bearing a median row of enlarged scales, as do many other karst-dwelling species. Material examined. Malaysia: Perlis: Perlis State Park LSUHC 8846–49, 8990–95, 9035. Thailand: Songkhla Province, Had Yai Disrict, Nga Chang Waterfall near the Ton Nga Waterfall MS 393–94.

Northern Sunda clade

The Northern Sunda clade is a lineage containing 28 species within four species groups (the chanthaburiensis, siamensis, argus, and affinis groups) that collectively frame the northern and western borders of the South China Sea from southern Vietnam to central Peninsular Malaysia (Figs. 2,3).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 39 75 Chanthaburiensis group. The chanthaburiensis species group is composed of six species that range across southern Indochina from southern Vietnam to Thailand along the mountainous coastline bordering the northern shores of the Gulf of Thailand. The basal lineage of this group is composed of the sister species Cnemaspis chanthaburiensis Bauer & Das from eastern Thailand and southwestern Cambodia and C. neangthyi Grismer, Grismer & Chav from southwestern Cambodia (Fig. 3). The sister lineage of this group is composed of the geographically proximate, microendemic, granite-dwellers C. aurantiacopes Grismer & Ngo; C. caudanivea Grismer & Ngo; C. nuicamensis Grismer & Ngo; and C. tucdupensis Grismer & Ngo from continental and insular southern Vietnam (Fig. 3). This is a well-supported lineage that is further supported here in that they are the only species of Cnemaspis that have a dark, mid-gular line, which we hypothesize to be a synapomorphy based on its absence from all other Cnemaspis and outgroups and thus constitutes further evidence of their monophyly. Their close, circumscribed, geographic proximity across a previously connected range of mountain tops (Grismer & Ngo 2007) is consistent with this hypothesis. This group is diagnosed by having a maximum SVL of 40.9–58.4 mm; 7–13 supralabials; 7–12 infralabials; smooth ventral scales; 0–9 pore-bearing precloacal pores; 16–31 paravertebral tubercles; smooth subcaudals bearing a slightly enlarged to enlarged median row; 0–4 postcloacal tubercles; no enlarged femoral or subtibial scales; and 22–33 subdigital lamellae on the fourth toe.

FIGURE 15. Cnemaspis chanthaburiensis from the Phnom Samkos Wildlife Sanctuary, Pursat Province, Cardamom Mountains, Cambodia. Upper left and right: adult males (LSUDPC 5056 and 5054, respectively) from Phnom Dalai in the dark and light color pattern phases, respectively. Lower left: subadult male (LSUDPC 2875) from the base of Phnom Samkos. Photographs by LLG. Lower right: ventral view of adult male (LSUDPC 8539) from Khao Kitchakut, Chanthaburi Province, Thailand. Photograph by PLW.

Cnemaspis chanthaburiensis Bauer & Das, 1998 Chanthaburi Rock Gecko Fig. 15

Cnemaspis sp. A Dring 1979:220

40 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 76 Holotype. FMNH 215979. Type locality: “Khao Soi Daow (Dao) Wildlife Sanctuary, Pongnomron (Pong Nam Ron), Chanthaburi Province, Thailand (approximately) 13°00’N, 102°05’E)” at 100 m in elevation. Diagnosis. Maximum SVL 42.2 mm; 8–10 supralabials; 7–10 infralabials; smooth ventral scales; 6–9 contiguous pore-bearing precloacal scales with elongate or round pores; 21–25 paravertebral tubercles; tubercles linearly arranged; caudal tubercles not restricted to a single paravertebral row nor encircling tail; tubercles in lateral caudal furrows; lateral row of tubercles present; subcaudals smooth, median row of scales ranging from not enlarged to weakly enlarged or only weakly enlarged posteriorly only; 1–3 postcloacal tubercles on each side; no enlarged femoral, subtibial or submetatarsal scales on first toe; subtibials smooth to weakly keeled; 22–29 subdigital fourth toe lamellae; in males gular region, throat, abdomen, and subcaudal region orange (Tables 6,7). Color pattern (Fig. 15). Ground color of top of head and rostrum yellowish; ground color yellowish dorsally on body, grayish on flank transitioning to orangish on tail; dorsum bearing alternating distinct light and smaller indistinct dark vertebral and paravertebral markings; light vertebral markings sometimes fused to form an irregularly shaped stripe; light markings form yellowish caudal bands; limbs bearing cream colored bands alternating with small, irregularly shaped black markings; chin, abdomen, and subcaudal region usually orange. Distribution. Cnemaspis chanthaburiensis is known from Khao Soi Dao Wildlife Sanctuary, Pong Nam Ron District, Chanthaburi Province; Khao Khieo Wildlife Sanctuary, Chon Buri Province; Khao Kitchakoot National Park, Chanthaburi Province; Khao Wong, Rayong Province; and Suan Kaset, Muang District and Namtok Pliieu, Chanthaburi Province in southeastern Thailand and from the base of Phnom Samkos, Phnom Dalai, and O’Som in the Samkos Wildlife Sanctuary of the Cardamom Mountains in southwestern Cambodia (Bauer & Das 1998; Grismer et al. 2008c,d; Fig. 3).

FIGURE 16. Habitat of Cnemaspis chanthaburiensis at Phnom Dalai, Phnom Samkos Wildlife Sanctuary, Pursat Province, Cardamom Mountains, Cambodia. Photograph by LLG.

Natural history. Cnemaspis chanthaburiensis is a small, secretive species restricted to hilly primary forests (Fig. 16) and ranges from near sea level to at least 968 m in elevation (Grismer et al. 2008d; Neang et al. 2010). Unlike most other species of Cnemaspis, C. chanthaburiensis is terrestrial and lizards are commonly found during the day inactive beneath and within logs and beneath loose bark on the forest floor (Neang et al. 2010). On two

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 41 77 occasions we have even collected specimens from the fly covers of our tents (Grismer et al. 2008d). We have found gravid females carrying two eggs during August. Relationships. Cnemaspis chanthaburiensis is the basal lineage of the chanthaburiensis group (Fig. 2). Material examined. Thailand: Chanthaburi Province, Pongnomron (Pong Nam Ron) District, Khao Soi Daouw (Dao) Wildlife Sanctuary, FMNH 215979 (holotype) and FMNH 191479 (paratype); Chantaburi Province, BMNH 1917.5.14.4 (paratype); Chon Buri Province, Khao Khiew (Khieo) Wildlife Sanctuary, FMNH 215978 (paratype); Chantaburi Province, Suan Kaset, Muang District, FMNH 215980 (paratype). Material examined since Grismer et al. (2010): Cambodia: Phnom Samkos, Pursat Province, Cardamom Mountains LSUHC 7882, Phnom Dalai 9337–38, near O’Som LSUHC 10110–11. Thailand: Chantaburi Province, Khao Khitchakut NP LSUHC 9507–08.

Cnemaspis neangthyi Grismer, Grismer & Thou, 2010 Neang Thy’s Rock Gecko Fig. 17

Holotype. LSUHC 8485. Type locality: “…outside the village of O’lakmeas, Pursat Province, Cambodia (12°19.4339N, 103°30.6059E)” at 145 m in elevation. Diagnosis. Maximum SVL 54.0 mm; 11–13 supralabials; 10–12 infralabials; smooth ventral scales; two round, contiguous pore-bearing, precloacal scales with round pores; 20–26 paravertebral tubercles; dorsal tubercles linearly arranged; caudal tubercles not restricted to a single paravertebral row; tubercles in lateral caudal furrows; lateral row of caudal tubercles present; ventrolateral caudal tubercles present anteriorly; caudal tubercles do not encircle tail; subcaudals smooth, bearing a median row of enlarged scales; one postcloacal tubercle on each side; no enlarged femoral, subtibial or submetatarsal scales on first toe; subtibials keeled; 22–25 subdigital fourth toe lamellae; lacks the diagnostic color pattern characteristics of the other species in the Indochina clade (Tables 6,7).

FIGURE 17. Upper left: adult male Cnemaspis neangthyi from near the village of O’lakmeas, Pursat Province, Cambodia. Right: siltstone microhabitat of C. neangthyi. Lower left: lowland, dry, deciduous dipterocarp forest habitat through which C. neangthyi ranges in isolated microhabitat islands. Photographs by LLG.

42 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 78 Color pattern (Fig. 17). Dorsal ground color of body and limbs olive-green to dull-yellow; head bearing a distinct, black parietal spot; radiating black and dull-white postorbital lines; light-green chevron marking on anterior margin of shoulder region; body overlain with light colored, paired, paravertebral blotches containing a central black dot and alternating with round, black blotches that extend onto tail to form poorly defined alternating dark and light bands; regenerated tail dull yellow, nearly unicolor; black spots on flanks invade lateral margins of abdomen; limbs stippled with light green and black; venter dull yellow with black stippling in scales. Distribution. Cnemaspis neangthyi is known only from the type locality at O'Lakmeas, Pursat Province, Cambodia (Fig. 3). Natural history. According to J. Grismer et al. (2010), Cnemaspis neangthyi is strictly nocturnal and found exclusively on boulders and cliff faces formed from sedimentary limestone-like rock. Cnemaspis neangthyi is most commonly found on vertical and overhanging faces, within cracks and shallow wind-eroded holes, shallow caves, and beneath exfoliations (Fig. 17). Cnemaspis neangthyi is adept at substrate matching and J. Grismer et al. (2010) found lizards only on sections of the rock wall containing light and dark green-colored lichens where their mottled color pattern enhanced their crypsis. No specimens were observed on the ground or in vegetation but when disturbed, lizards would run to the base of the rock wall to hide in cracks and holes formed by the expansive soil at the ground-rock interface. Relationships. Cnemaspis neangthyi is the sister species of the Vietnamese lineage containing C. aurantiacopes, C. caudanivea, C. tucdupensis, and C. nuicamensis (Fig. 2). Material examined. Cambodia: Pursat Province, O’Lakmeas LSUHC 8478, 8485, 8515–17 (type series).

Cnemaspis aurantiacopes Grismer & Ngo, 2007 Hon Dat Rock Gecko Fig. 18

Holotype. UNS 49. Type locality: Hon Dat Hill, Hon Dat District, Kien Giang Province, Vietnam (10°06.7749 N, 104°53.5699 E)” at 30 m in elevation. Diagnosis. Maximum SVL 58.4 mm; 9–11 supralabials; 8–10 infralabials; smooth ventral scales; no precloacal pores; 23–31 paravertebral tubercles; tubercles on body linearly arranged and present on flanks; caudal tubercles not restricted to a single paravertebral row; no tubercles in lateral caudal furrows; ventrolateral caudal tubercles present anteriorly; caudal tubercles do not encircle tail; lateral caudal tubercle row present; subcaudals smooth, bearing a median row of enlarged scales; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; slightly enlarged submetatarsal scales on first toe; 27–31 subdigital fourth toe lamellae; faint, dark, elongate mid-gular marking; gukar region, throat, pectoral region, dorsal and ventral surfaces of forelimbs, ventral surface of hind limbs, and original tail orange in males (Tables 6,7). Color pattern (Fig. 18). Males: dorsal ground color yellowish to saffron, overlain by rust colored, semi- transversely oriented, irregularly shaped markings extending from occiput to base of tail and enclosing a series of eight, large, yellowish-gray, oval blotches extending from nape to base of tail; top of head reddish-brown, rostrum gray; dorsal surface of limbs saffron, overlain with faint, lighter mottling on brachia and thighs and weak banding on forelimbs and forelegs; dorsal caudal region reddish-brown, no banding; three wide, faint, reddish brown, postorbital stripes, uppermost extending onto shoulder region and contacting blotch on nape; ventral surfaces of neck, body, and limbs dull orange, immaculate; gular region slightly darker; labials unicolor reddish brown. Females: overall dorsal coloration is more yellowish-gray, especially noticeable on the limbs; dark, rhomboid blotches and bands on all dorsal surfaces. Both sexes bear a faint, usually lineate, median mid-gular marking. Distribution. Cnemaspis auarntiacopes is known only from the type locality at Hon Dat Hill, Hon Dat District, Kien Giang Province, Vietnam (Fig. 3). Natural history. Hon Dat Hill composes a small cluster of mountains reaching to 100 m in elevation that supports an isolated section of secondary, highly disturbed, semi-deciduous forest in the southern reaches of the Mekong Delta flood plain (Fig. 3) and is completely surrounded by agricultural lowlands. The hill maintains abundant outcroppings of granitic rocks and caves and Grismer & Ngo (2007) noted Cnemaspis aurantiacopes to be common on outcrops in both highly disturbed and old secondary forest. They did not observe lizards during the day but lizards were common at night within the confines of caves. This species does not venture out onto the open

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 43 79 surfaces of the boulders and when alarmed, may give a tail display wherein the tail is rolled slightly over the back and often moved from side to side (Grismer & Ngo 2007). Relationships. Cnemaspis aurantiacopes is the basal species of the other Vietnamese species of Cnemaspis; C. caudanivea Grismer & Ngo, C. tucdupensis Grismer & Ngo, and C. nuicamensis Grismer & Ngo (Fig. 2). Material examined. Vietnam: Kien Giang Province, Hon Dat District, Hon Dat Hill UNS 47, 49 (type series). Additional material examined since Grismer & Ngo (2007): LSUHC 8245, 9528–41.

FIGURE 18. Cnemaspis aurantiacopes from Hon Dat Hill, Kien Giang Province, Vietnam. Upper and lower left: adult males (LSUDPC 3131 and 3130, respectively) in the light color pattern phase. Upper right: subadult male (LSUDPC 3135) in the light color pattern phase. Lower right: adult female (LSUDPC 3133) in the dark color pattern phase. Photographs by LLG.

Cnemaspis caudanivea Grismer & Ngo, 2007 Hon Tre Island Rock Gecko Fig. 19

Holotype. UNS 83. Type locality: “Hon Tre Island, Kien Hai District, Kien Giang Province, Vietnam (09°58.3429 N, 104°50.9559 E)” at 100 m in elevation. Diagnosis. Maximum SVL 47.2 mm; eight or nine supralabials; seven or eight infralabials; smooth ventral scales; 0–2 round, discontinuous, pore-bearing, precloacal scales; 20–24 paravertebral tubercles; tubercles linearly arranged but absent from flanks; caudal tubercles not restricted to a single paravertebral row nor encircling tail; caudal tubercles occasionally in lateral caudal furrows anteriorly only; ventrolateral caudal tubercles variably present anteriorly; lateral caudal tubercle row occasionally present anteriorly; subcaudals smooth, sometimes bearing a median row of slightly enlarged scales; one or two postcloacal tubercles on each side; no enlarged femoral or subtibial scales; subtibials smooth to weakly keeled; weakly enlarged submetatarsal scales occasionally present on first toe; 23–30 subdigital fourth toe lamellae; large, black, round spots on nape and anterior portion of body; dark elongate mid-gular marking; wide black and yellow bands on tail; posterior portion of tail immaculate white dorsally and ventrally (Tables 6,7).

44 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 80 FIGURE 19. Cnemaspis caudanivea from Hon Tre Island, Kien Giang Province, Vietnam. Upper left: adult male (LSUDPC 3056) in the light color pattern phase. Upper right: adult female (LSUDPC 3045) in the dark color pattern phase. Lower: granite boulder microhabitat on Hon Tre Island. Photographs by LLG.

Color pattern (Fig. 19). Dorsal ground color gray, overlain by a dark pattern of reddish brown, irregularly shaped markings on top of head and snout; squarish, semi-transversely arranged, large, black blotches on neck and body separated by dull white blotches; irregularly shaped bands on limbs; wide, black and dull-yellow bands encircling tail; wide, poorly defined, dark-brown postorbital stripes edged below in white extending onto side of neck; ventral surfaces of neck, body, and limbs dull beige, immaculate; gular region smudged with darker coloration and light spots, bearing a dark mid-gular stripe; last 25% of original tail immaculate, brilliant white dorsally and ventrally. Distribution. Cnemaspis caudanivea is known only from the type locality of Hon Tre Island, Kien Hai District, Kien Giang Province, Vietnam (Fig. 3). Natural history. Hon Tre is a small (70 km2) island lying 15 km off the southern coast of Vietnam in Rach Gia Bay, 28.3 km nearly due west of the port of Rach Gia in Kien Giang Province. The island is steep-sided, reaching 315 m in elevation and dominated by secondary and primary semi-deciduous forest covering a landscape composed of nearly continuous outcrops of large, granitic boulders (Fig. 19). Grismer & Ngo (2007) noted lizards occurred on granite boulders throughout the island, from the coastline to near the summit. During the day, lizards

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 45 81 are only observed on shady, inclined surfaces and within rock cracks and cave-like cavities formed by boulders piled on top of one another. At night, lizards venture onto the exposed outer surfaces of the boulders but appear inactive. Lizards are exceptionally wary both day and night and typically escape capture at surprisingly high speeds, often hopping while running. Grismer & Ngo (2007) reported lizards curling and elevating their tail above their back when alarmed and moving the posterior 25% from side to side, thus displaying the strikingly white coloration. The dexterity with which the tail is waved back and forth, however, exceeds that of all other Cnemaspis we have observed. Cnemaspis caudanivea can fold the end of the tail forward 180° on itself while rolling and unrolling it slowly in a lateral plane, making it look like a small, white worm wiggling on a rock. This behavior and color pattern is maintained in regenerated tails as well, suggesting it is strongly selected for. Relationships. Cnemaspis caudanivea is the sister species to a monophyletic group composed of C. tucdupensis and C. nuicamensis (Fig. 2). Material examined. Vietnam: Kien Giang Province, Kien Hai District, Hon Tre Island UNS 83, 84 (type series). Material examined since Grismer & Ngo (2007): Vietnam; Kien Giang Province, Kien Hai District, Hon Tre Island LSUHC 8247, 9543–48.

Cnemaspis nuicamensis Grismer & Ngo, 2007 Nui Cam Hill Rock Gecko Fig. 20

Holotype. UNS 37. Type locality: “Nui Cam Hill, Tinh Bien District, An Giang Province, Vietnam (10°29.7759 N, 105°00.4419 E)” at 100 m in elevation. Diagnosis. Maximum SVL 48.2 mm; 7–9 supralabials; six or seven infralabials; smooth ventral scales; 3–6 pore-bearing, precloacal scales separated medially by 1–4 poreless scales; 16–21 paravertebral tubercles; tubercles linearly arranged and present on flanks; caudal tubercles not restricted to a single paravertebral row nor encircling tail; tubercles in lateral caudal furrows occasionally anteriorly; ventrolateral caudal tubercles present anteriorly; lateral caudal tubercle row absent or present only anteriorly; subcaudals smooth and bearing a median row of enlarged scales ; 2–4 postcloacal tubercles on each side; no enlarged femoral or subtibial scales; subtibials smooth; no enlarged submetatarsal scales on first toe; 27–33 subdigital fourth toe lamellae; dark, elongate mid-gular marking (Tables 6,7). Color pattern (Fig. 20). Dorsal ground color dull-yellow, body overlain with large, reddish brown blotches and offset, black, paravertebral blotches; limbs with reddish brown and pale yellow, alternating bands; poorly defined, reddish brown, caudal bands not encircling original tail; top of head and snout with reddish brown reticulum enclosing dull white blotches; scattered, dark spots on top of head; three thin, dark, reddish brown, postorbital stripes infused with black and edged below in white, uppermost not extending onto shoulder region; all ventral surfaces gray, immaculate; gular region bearing a dark, mid-gular line. Distribution. Cnemaspis nuicamensis is known only from the type locality of Nui Cam Hill 22.5 km south of Chau Doc in Tinh Bien District of An Giang Province, Vietnam near the border of Cambodia (Fig. 3). Natural history. Nui Cam Hill is part of the Bay Nui Mountains, which are surrounded by the vast, agricultural lowlands of the Mekong Delta flood plain. Nui Cam Hill is the tallest (710 m) in this range of mountains and is dominated by old growth, secondary, semi-deciduous forest and highly disturbed forest. Lizards were all observed during the day on the surface of granite rocks or within rock cracks. Within this microhabitat, Cnemaspis nuicamensis is far more common on vertical surfaces in the vicinity of streams than on rocks within the forest (Fig. 20). Grismer & Ngo (2007) noted that lizards presented tail displays upon escape and no lizards were observed on rocks within disturbed forest. Relationships. Cnemaspis nuicamensis is the sister species to C. tucdupensis (Fig. 2). Material examined. Vietnam: An Giang Province, Tinh Bien District, Nui Cam Hill UNS 37, 38 (type series). Material examined since Grismer & Ngo (2007): Vietnam; An Giang Province, Tinh Bien District, Nui Cam Hill LSUHC 8246, 8646–52, 9549–55.

46 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 82 FIGURE 20. Upper; adult male Cnemaspis nuicamensis (LSUDPC 3136) in the light color pattern phase. Lower: Granite microhabitat of C. nuicamensis at Nui Cam Hill, An Giang Province, Vietnam. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 47 83 Cnemaspis tucdupensis Grismer & Ngo, 2007 Tuc Dup Hill Rock Gecko Fig. 21

Holotype. UNS 45. Type locality: “Tuc Dup Hill, Tri Ton District, An Giang Province, Vietnam (10°22.9549 N, 104°57.3369 E) at 100 m in elevation. Diagnosis. Maximum SVL 51.0 mm; 8–10 supralabials; 7–9 infralabials; smooth ventral scales; no precloacal pores; 16–22 paravertebral tubercles; tubercles linearly arranged or nearly so, present on flanks; caudal tubercles not restricted to a single paravertebral row nor encircling tail; tubercles not in lateral caudal furrows; ventrolateral caudal tubercles present anteriorly; no lateral caudal tubercle row; subcaudals smooth, bearing a median row of slightly enlarged scales; 0–3 postcloacal tubercles on each side; no enlarged femoral or subtibial scales; subtibials smooth; enlarged submetatarsal scales on first toe; 26–32 subdigital fourth toe lamellae; large, black, squarish, paired paravertebral markings on body; black and yellow bands on tail; posterior portion of original tail in males black; dark mid-gular marking; gular region, throat, pectoral region, abdomen, ventral surfaces of limbs and subcaudal region in males orangish (Tables 6,7).

FIGURE 21. Cnemaspis tucdupensis from Tuc Dup Hill, An Giang Province, Vietnam. Upper left: adult female (LSUDPC 3140) in the light color pattern phase. Upper right: adult male (LSUDCP 3141) in the dark color pattern phase. Lower: granite boulder microhabitat at Tuc Dup Hill. Photographs by LLG.

Color pattern (Fig. 21). Dorsal ground color gray, body overlain with large, black and pale yellow spots; limbs bearing reddish brown and pale yellow alternating bands; poorly defined, black and yellow caudal bands encircling tail; posterior portion of original tail black; regenerated tail beige, immaculate; top of head and snout with faint, dark reticulum enclosing dull whitish blotches; thin, dark, postorbital stripes edged below in white,

48 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 84 uppermost extending onto nape; dark mid-gular marking; gular region, throat, pectoral region, abdomen, ventral surfaces of limbs and subcaudal region in males orangish. Distribution. Cnemaspis tucdupensis is known only from the type locality of Tuc Dup Hill located 36.3 km south of Chau Doc in Tinh Bien District of An Giang Province, Vietnam near the border of Cambodia (Fig. 3). Natural history. Tuc Dup Hill is a prominence of Co To Mountain that lies at the southern end of the Bay Nui Mountains in Tri Ton District of An Giang Province in southern Vietnam. Co To Mountain reaches 584 m in elevation and is covered with primary semi-deciduous forest where numerous granitic outcroppings and caves are common (Fig. 21). At Tuc Dup, the boulders are particularly large (up to 5 m in diameter) and enclose a myriad of spaces and chambers where they lie on top of one another. The type locality served as a Viet Cong headquarters during the Vietnam War and currently serves as a museum celebrating Tuc Dup Hill as a strategic military hideout. Cnemaspis tucdupensis is most prevalent in caverns near areas wherein sunlight can filter down during the day supporting plant growth and mosses on the boulders and serving as areas where lizards can forage (Grismer & Ngo 2007). Lizards avoid direct sunlight and remain on the shaded vertical and inverted rocky surfaces nearby. At night, lizards venture out into open areas within the cavern as well as onto boulder surfaces exposed to the outside and appear inactive. Lizards present weak tail displays immediately before and after escaping. Relationships. Cnemaspis tucdupensis is the sister species of C. nuicamensis (Fig. 2). Material examined. Vietnam: An Giang Province, Tri Ton District, Tuc Dup Hill UNS 42–43, 45 (type series). Material examined since Grismer & Ngo (2007): Vietnam; An Giang Province, Tri Ton District, Tuc Dup Hill LSUHC 8245, 8609–22, 9527–41. Grismer & Ngo (2007) mistakenly listed the holotype as UNS 49 when it should have been UNS 45. Siamensis group. The siamensis group is a well-supported lineage that contains parapatric sister lineages on opposite sides of the Isthmus of Kra in the central portion of the Thai-Malay Peninsula (Figs. 2,3). This group may be the sister lineage to the chanthaburiensis group although this relationship lacks strong statistical support (Fig. 2). The northern lineage contains C. siamensis (Smith) that extends throughout northern Peninsular Thailand (and probably Myanmar) and its sister species C. huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya known only from western Thailand just north of the Thai-Malay Peninsula (Grismer et al. 2010a). The southern lineage contains C. chanardi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya in the north and the sister species C. omari sp. nov. from southern Thailand and northern Peninsular Malaysia and C. roticanai Grismer & Chan from Langkawi Island in extreme northwestern Malaysia (Fig. 3). These three species form a well-supported monophyletic group in the molecular analysis that is further supported here by them having the derived character states of a light-colored prescapular crescent, a yellow belly, and yellow ventral surfaces of the hind limbs. Some individuals of C. biocellata from the Pattani clade have a yellow belly, which we consider convergent. The siamensis group is diagnosed by having a maximum SVL of 37.8–47.0 mm; 7–10 supralabials; 6–9 infralabials; 0–8 pore-bearing, precloacal scales; linearly arranged dorsal tubercles; 18–30 paravertebral tubercles; caudal tubercles not restricted to a single paravertebral row and encircling tail; no ventrolateral caudal tubercles anteriorly; one or two postcloacal tubercles on each side; no enlarged femoral, subtibial or scales beneath first metatarsal; 21–31 lamellae beneath the fourth toe.

Cnemaspis siamensis (Smith, 1925) Siam Rock Gecko Fig. 22

Gonatodes siamensis Smith, M. A., 1925:21. Gonatodes kendallii Smith, M. A. 1916:151. non Boulenger fide Taylor, 1963:740. Gonatodes siamensis Smith, 1930:16.

Holotype. BMNH 1946.8.19.83. Type locality: “Maprit, near Patiyu (=Pathio, Chumpon Province), Peninsular Siam [Thailand]” at 10 m in elevation. Diagnosis. Maximum SVL 39.7 mm; eight or nine supralabials; 6–8 infralabials; keeled ventral scales; no precloacal pores; 19–25 paravertebral tubercles; dorsal tubercles on body randomly arranged; tubercles on flanks;

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 49 85 caudal tubercles not restricted to a single paravertebral row nor encircling tail; no tubercles in lateral caudal furrows; no ventrolateral caudal tubercles; lateral caudal tubercle row present; subcaudals keeled; median row of enlarged subcaudal scales present; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 24–26 subdigital fourth toe lamellae; gular region, throat, and pectoral region yellow in males; dark, longitudinally arranged spots in gular region (Tables 6,7). Color pattern (Fig. 22). Dorsal ground color gray to light brown overlain with short, dark, zig-zag, transverse bands on body countershaded with white markings; top of head and flanks mottled with light and dark irregularly shaped markings; limbs and tail bearing poorly defined dark bands; gular region, chest and posterior portion of original tail in males yellow; dark longitudinally arranged spots on a yellow gular region; belly and anterior portion of tail cream colored, immaculate; ventral surfaces of limbs gray.

FIGURE 22. Cnemaspis siamensis from Pathio, Chumpon Province, Thailand. Upper right: adult male (LSUDPC 5241) in the dark color pattern phase. Middle right: adult female (LSUDPC 5228) in the light color pattern phase. Lower right: ventral coloration of adult male (LSUDPC 5242). Left: microhabitat structure. Photographs by LLG.

Distribution. Cnemaspis siamensis ranges throughout the lowland, hilly regions east of the Tenasserim and Phuket Mountains from Kaeng Krachan National Park, Phetchaburi Province in the north, southward to Khao Mod, Surat Thani Province on the east coast and to Phuket Island, Phuket Province in the west (Grismer et al. 2010a; Fig. 3).

50 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 86 Natural history. Grismer et al. (2010a) stated that Cnemaspis siamensis is not a saxicolous, microhabitat specialist but a nocturnal, lowland, scansorial, forest-dwelling gecko that opportunistically utilizes rocky microhabitats when available. We have observed lizards at the type locality on both vegetation (Fig. 22) and rocks. Relationships. Cnemaspis siamensis is the sister species of C. huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya (Fig. 2). Material examined. Thailand: Chumphon Province, Krom Luang District THNHM 0372; Pha To District, Ngao National Park THNHM 1086; Pathio LSUHC 9474, 9485, MCZ 39025; Kapoh Water Fall FMNH 215977. Phetchaburi Province, Muang District THNHM 1441–42, 1448–49. Prachuap Khiri Khan Province, Pa-La-U, Kaeng Krachan National Park, Hua Hin District, Prachuap Khiri Khan THNHM 1336–37; Thap Sakae District THNHM 2000. Surat Thani Province, Kanchanadit District MS16, Kaeng Krung National Park THNHM 1084.

FIGURE 23. Cnemaspis huaseesom from Sai Yok National Park, Kanchanaburi Province, Thailand. Upper left; ventral views showing color pattern variation in adult males (LSUDPC 5225). Upper right; adult male (LSUDPC 5213) in the dark color pattern phase. Lower right: adult female (LSUDPC 5214) in the light color pattern phase. Photographs by LLG. Lower left: adult male (LSUDPC 8276) showing variation in color pattern in that it lacks a yellow tail. Photograph by MS.

Cnemaspis huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Yellow-headed Rock Gecko Figs. 23,24

Holotype. THNHM 15909. Type locality: “Sai Yok National Park, Kanchanaburi Province, Thailand (14°20.09N, 98°51.35E)” at 125 m in elevation. Diagnosis. Maximum SVL 43.5 mm; 7–10 supralabials; 6–9 infralabials; smooth ventral scales; 5–8 contiguous, pore-bearing, precloacal scales with round pores; 18–24 paravertebral tubercles; tubercles on flanks; tubercles in lateral caudal furrows; no ventrolateral caudal tubercles; lateral caudal tubercle row absent; subcaudals smooth, with no enlarged or weakly keeled median scale row; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials smooth; no enlarged submetatarsal scales on first toe; 21–31 subdigital fourth toe lamellae; head, forelimbs, tail, gular region, throat, pectoral region, underside of forelimbs, and subcaudal region yellow in males (Tables 6,7). Color pattern (Figs. 23,24). Males: Cnemaspis huaseesom are capable of considerable change in coloration from a light to darker phase. The description that follows is of the darker phase: dorsal ground color of head,

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 51 87 forelimbs and tail yellow, with faint banding on tail; dorsal ground color of trunk and hind limbs dark-gray and bearing large, light-gray, paravertebral spots extending from occiput to base of tail; dorsal surface of hind limbs bearing large, light-gray spots; dorsal surface of forelimbs bearing small, yellow markings; top of head mottled; dark postorbital stripes faint; large, round, whitish markings on nape; trunk uniformly gray; tubercles on body lightly colored; belly pale gray; ventral surface of hind limbs gray; fine, dark stippling on all ventral surfaces, most dense on belly. Some adult males may have a gray, as opposed to yellow tail. Females: lack yellow head, forelimbs, and tail, have same general trunk color as males in dark phase; overall ground color of head, body, limbs, and tail light brown; large, lighter, paravertebral markings extend from nape to base of tail where they continue posteriorly to form lightly colored, caudal bands; flanks densely stippled with cream-colored markings and bear faint, gray bars; limbs mottled; all ventral surfaces beige with faint stippling that is most dense on belly and tail. Distribution. Cnemaspis huaseesom is known only from the type locality of Sai Yok National Park, Kanchanaburi Province, Thailand (Fig. 3). Natural history. Grismer et al. (2010a) stated that Cnemaspis huaseesom is most commonly found on hillsides in lowland areas with karst boulders in semideciduous, dipterocarp forest amongst thick vegetation including bamboo. Lizards are generally active at night on karst boulders but may be found on vine-like vegetation near the boulders (Fig. 24). This species is fast, wary, and flees into deep cracks and crevices at the slightest provocation. More than one lizard is usually found on a given outcropping. Lizards are only rarely observed during the day. Relationships. Cnemaspis huaseesom is the sister species of C. siamensis (Fig. 2). Material examined. Thailand: Kanchanaburi Province, Sai Yok National Park THNHM 15909, PSUZC-RT 2010.55, CUMZ-R 2009, 6, 24–4 (type series). Material examined since Grismer et al. (2010a): Thailand: Kanchanaburi Province, Sai Yok National Park LSUHC 9455–58.

FIGURE 24. Left: karst microhabitat of Cnemaspis huaseesom at Sai Yok National Park, Kanchanaburi Province, Thailand. Photograph by LLG. Right: adult female (LSUDPC 8278) from Sai Yok National Park in the light color pattern phase. Photograph by MS.

Cnemaspis chanardi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Chan-ard’s Rock gecko Fig. 25

Gonatodes siamensis Smith, 1930:16 (in part) Cnemaspis siamensis Smith, 1935:71; Taylor, 1963:740 (in part)

52 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 88 Holotype. THNHM 6983. Type locality: “Ban Chong, Chong, Nayong District, Trang Province, Thailand…between 400 and 600 m a.s.l.” Diagnosis. Maximum SVL 40.1 mm; 7–10 supralabials; 6–8 infralabials; keeled ventral scales; 6–8 discontinuous pore-bearing precloacal scales with round pores; 20–30 paravertebral tubercles; tubercles on flanks; no tubercles in lateral caudal furrows; ventrolateral caudal tubercles absent anteriorly; caudal tubercles not encircling tail; lateral caudal tubercle row present; subcaudals keeled; median row of enlarged subcaudal scales present; one postcloacal tubercle on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 25–30 subdigital fourth toe lamellae; yellowish, prescapular crescent; gular region, belly, underside of hind limbs, and subcaudal region yellow in males (Tables 6,7). Color pattern (Fig. 25). Dorsal ground color of head, body, limbs and tail gray; top of head bearing small, diffuse, faint, darker colored markings giving it a somewhat mottled appearance; dark postorbital stripes faint; large, round, whitish markings on nape; trunk uniformly gray except for lightly colored tubercles on body; forelimbs mottled bearing a slight banding pattern; ventral surface of gular region, abdomen, hind limbs and tail yellow; abdomen and hind limbs sometimes pale gray; fine, dark stippling on all ventral surfaces, most dense on belly.

FIGURE 25. Left: tree hole microhabitat of Cnemaspis chanardi at the Phuphaphet Cave area, Satun Province, Thailand. Upper right: adult male C. chanardi (LSUDPC 8304) from Dad Fa Waterfall, Surat Thani Province, Thailand in the dark color pattern phase. Middle right: adult female (LSUDPC 8288) from Khao Luang National Park, Nakhon Si Thammarat Province, Thailand in the dark color pattern phase. Lower right: ventral view of adult male (LSUDPC 8301) from Dad Fa Waterfall, Surat Thani Province, Thailand. Photographs by MS.

Distribution. Cnemaspis chanardi occurs in the foothills of the Nakhon Si Thammarat and Sankalakhiri Mountains and lowland regions from the southern terminus of the Isthmus of Kra in Donsak District, Surat Thani Province, southward to Khao Chong and Nayong district, Trang Province. From the foothills of these mountains, C. chanardi extends westward through the lowlands to at least Khlong Thom District, Krabi Province (Fig. 3). Cnemaspis chanardi is not known to occur in the foothills or lowlands east of the crests of the Nakhon Si Thammarat and Sankalakhiri mountains (Fig. 3). Grismer et al. (2010a) considered the presence of C. chanardi (as opposed to the geographically more proximate C. siamensis) on Ko Tao Island approximately 85 km off the coast

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 53 89 from Muang District, Chumpon Province, (Fig. 3), consistent with the geological history of this part of Peninsular Thailand. The island chain consisting of Ko Tao and the intervening islands Ko Samui and Ko Phangan, are offshore extensions of the Nakhon Si Thammarat Mountains to which they were connected during the last glacial maximum (Sathiamurthy & Voris 2006) and lie to the east of the Isthmus of Kra (Fig. 3). It is likely that C. chanardi also occurs on Ko Samui and Ko Phangan. Natural history. Grismer et al. (2010a) and Taylor (1963) noted that Cnemaspis chanardi is a diurnal, scansorial species that utilizes large, open, above-ground substrates (i.e. tree trunks and boulders; Fig. 25) and does not occupy habitats that do not contain both trees and rocks. Lizards are occasionally found beneath small rocks. Cnemaspis chanardi ranges from near sea level at Khlong Thom to just under 600 m at Khao Chong. Smith (1930) reported specimens from Khao Whip (=Khao Wang Hip), Nakhon Si Thammarat Province but gave no elevation. Remarks. Based on the geographically outlying southernmost paratype LSUHC 9564 from the Phuphaphet Cave, Satun Province, Thailand, Grismer et al. (2010a) considered this locality to be southern extent of the known range of Cnemaspis chanardi but posited that its range probably continued approximately 45 km further south to the Banjaran Nakawan mountains on the Thai-Malaysian border. However, genetic data from LSUHC 9565 from the same cave and from LSUHC 9978–79 from the Banjaran Nakawan mountains in Perlis, Malaysia (considered to be C. roticanai by Grismer [2011a]) indicate these populations are conspecific, fall outside of C. chanardi and C. roticanai, and are most closely related to the latter. Therefore, we consider the Phuphaphet Cave and Perlis populations to be the new species C. omari sp. nov. described below. We remove LSUHC 9564 from the type series of C. chanardi and consider C. roticanai to be endemic to Langkawi Island as was originally considered (Grismer & Chan 2010; Fig. 3). Relationshsips. Cnemaspis chanardi is the sister species of the lineage containing C. omari sp. nov. and C. roticanai Grismer & Chan (Fig. 2). Material examined. Thailand: Trang Province, Nayong District, Ban Chong, Chong, FMNH 176863, THNHM 6983 (holotype); Surat Thani Province, Ban Nasan District, Dad Fa Waterfall in Tai Rom Yen National Park CUMZ-R-2009, 6, 24-6, Donsak District, near Donsak Pier MS 395; Kanchanadit District, Petphanomwung Cave ZMKU Rep-000313, KZM 009; Ko Tao Island USNM 76143–44; Krabi Province, Khlong Thom District, Khao Nor Chuchi, Khlong Thom THNHM 12434, 12439–40; Nakhon Si Thammarat Province, Lan Saka District, Khao Luang National Park THNHM 1334-35, 14111, Nopphitam District, Krung Nang Waterfall in Khao Nan National Park, THNHM 10705, Khao Nan National Park, THNHM 10115, Tha Sala District, Khao Nan National Park THNHM 10135, Nopphitham District, Ban Yod Leong, THNHM 10383, Thum Panra District; Thum Thong Panra, PSUZC-RT 2010.53–54.

Cnemaspis omari sp. nov. Omar’s Rock Gecko Fig. 26

Cnemaspis chanardi Grismer et al. 2010a:24 (in part) Cnemaspis roticanai Grismer 2011a:367 (in part)

Holotype. Adult male (LSUHC 9978) collected by Evan S. H. Quah and M. A. Muin on 11 March 2011 at Wang Kelian, Perlis, Peninsular Malaysia (06°41.805 N, 100°10.751 E) at 150 meters above sea level. Paratypes. Adult female (LSUHC 9979) bears the same data as the holotype. Adult females (LSUHC 9564–65) collected by Kirati Kunya from the Phuphaphet Cave, Muang District, Satun Province, Thailand on 6 October 2009. LSUHC 9564 was previously considered a paratype of C. chanardi (Grismer et al. 2010a:17). Diagnosis. Maximum SVL 41.3 mm; eight or nine supralabials; seven or eight infralabials; keeled ventral scales; four contiguous pore-bearing precloacal scales with round pores; body tuberculation strong; 22–29 paravertebral tubercles; dorsal tubercles bear weak linear arrangement; tubercles present on flanks; no tubercles in lateral caudal furrows; ventrolateral caudal tubercles absent; caudal tubercles encircling tail; lateral caudal tubercle row present; subcaudals keeled, no enlarged median row; one postcloacal tubercle on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; and 25–28 subdigital fourth toe lamellae; light colored prescapular crescent; gular region, belly, underside of hind limbs, and subcaudal region yellow in males (Tables 6,7).

54 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 90 FIGURE 26. Cnemaspis omari sp. nov. Upper left: adult male holotype (LSUHC 9978) from Perlis State Park, Perlis, Peninsular Malaysia in the light color pattern phase. Middle left: adult female paratype (LSUHC 9979) from Perlis State Park, Perlis, Peninsular Malaysia in the light color pattern phase. Photographs by ESHQ. Upper and middle right: adult female paratype (LSUHC 9564) and subadult (LSUDPC 8299), respectively, from Phuphapet Cave, Satun Province, Thailand in dark color pattern phase. Photographs by MS. Lower: type series of C. omari sp. nov. Photograph by LLG.

Description of holotype. Adult male; SVL 41.3 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.27), somewhat narrow (HW/SVL 0.17), flat (HD/HL 0.41), distinct from neck; snout short (ES/HL 0.50), concave in lateral profile; postnasal region constricted medially, raised; scales of rostrum weakly keeled, larger than similarly shaped scales on occiput; moderate, supraorbital ridges; shallow frontonasal sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.23); extra-brillar fringe scales small in general but largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 75% divided by longitudinal groove; rostral bordered posteriorly by two supranasals and one smaller azygous scale, laterally by first supralabials and nostrils; 8R,L raised supralabials of similar size; 7R,L infralabials, decreasing gradually in size posteriorly; nostrils small, oblong, oriented dorsoposteriorly, bordered posteriorly by small, granular, postnasal scales; mental large,

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 55 91 triangular, medially concave, extending to level of second infralabial, bordered posteriorly by three postmentals, lateral postmentals largest; gular and throat scales raised, smooth, somewhat pointed; throat scales larger. Body slender, elongate (AG/SVL 0.50); small, weakly keeled, dorsal scales equal in size throughout body, intermixed with numerous, large, multi-keeled, semi-longitudinally arranged tubercles; tubercles extend from occiput to base of tail and are smallest anteriorly; 29 paravertebral tubercles; pectoral and abdominal scales flat, keeled, subimbricate, equal in size, much larger than dorsals; four pore-bearing, precloacal scales arranged in a 2(R)–2(L) chevron, separated medially by two non-pore-bearing scales; forelimbs moderately long (FL/SVL 0.17), slender, dorsal scales keeled; ventral scales of forearm smooth, juxtaposed to subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; subdigital lamellae wide throughout proximal portion of digits to inflection, more granular after inflection, bearing a larger scale at the digital inflections; interdigital webbing absent; fingers increase in length from first to fourth with fifth slightly shorter than fourth; hind limbs longer and thicker than forelimbs (TBL/SVL 0.21); dorsal scales keeled, raised, juxtaposed; ventral scales of thigh, raised, keeled; subtibials keeled, larger than dorsal tibials; plantar scales smooth, slightly raised, juxtaposed; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout length of digits except at base where scales are more granular; enlarged, scale at the digital inflections; interdigital webbing absent; toes increase in length from first to fourth with fourth being longest; 27R,26L subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; caudal scales raised, keeled, juxtaposed anteriorly; moderate, middorsal furrow; deep, single, lateral furrow; subcaudals keeled, no enlarged median row of scales; opposing paravertebral, dorsolateral, and lateral rows of large, keeled, equally sized, caudal tubercles; no ventrolateral caudal tubercles; caudal tubercles encircle tail, absent from lateral furrow; 1R,L postcloacal tubercle on lateral surface of hemipenal swellings at base of tail; anterior 11.9 mm original, reamainder 17.0 mm regenerated. Coloration in life (Fig. 26). Dorsal ground color of head, body, limbs and tail pale-yellow; rostrum bearing diffuse, faint brown spots; single, diffuse, black, postorbital stripe extending to occiput; diffuse, light yellow, oblong, vertebral marking on nape; paired, brown nape spots followed by pair of brown, paravertebral spots on neck followed by five small, irregularly shaped, brown, spots terminating at base of tail; original portion of tail faintly banded; regenerated portion of tail uniform pale gray; butterfly-shaped, pale-yellow, vertebral markings between paired, brown spots; yellow prescapular crescent followed by faint, semi-transversely arranged, yellow bars on flanks; forelimbs bearing yellowish blotches and scattered faint, dark markings; hind limbs bearing yellowish blotches and dark markings resembling a reticulate pattern; digits bearing dark bands; gular region yellowish orange; throat beige; abdomen, ventral surface of hind limbs, and subcaudal region yellow; ventral surface of forelimbs beige; all ventral scales bearing small, black stippling. Variation. The paratypes resemble the holotype in coloration (Fig. 26). The difference is that the Thai specimens are generally darker and more boldly patterned. The light nape marking on LSUHC 9979 is particularly apparent. Meristic and mensural differences are presented in Table 8.

TABLE 8. Meristic and mensural character states of the type series of Cnemaspis omari sp. nov. w = weak; f = female; and m = male. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right.

LSUHC LSUHC LSUHC LSUHC 9564 9565 9978 9979 paratype paratype holotype paratype SVL (mm) 40.5 39.7 41.3 39.9 Supralabials 9 9 8 9 Infralabials 8 8 7 8 Ventral scales keeled (1) or not (0) 1 1 1 1 No. of precloacal pores / / 4 / Precloacal pores continuous (1) or separated (0) / / 0 / Precloacal pores elongate (1) or round (0) / / 0 / No. of paravertebral tubercles 22 26 29 24 ...... continued on the next page

56 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 92 TABLE 8. (Continued) LSUHC LSUHC LSUHC LSUHC 9564 9565 9978 9979 paratype paratype holotype paratype Tubercles linearly arranged (1) or more random (0) 0 0 w w Tubercles present (1) or absent (0) on flanks 1 w 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 / 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 / 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 / 1 1 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 / 0 0 Subcaudals keeled (1) or not (0) 0 / 1 1 Single median row of keeled subcaudals (1) or not (0) 0 / 0 0 Caudal tubercles encircle tail (1) or not (0) 1 / 1 1 Enlarged median subcaudal scale row (1) or not (0) 0 / 0 0 Postcloacal spurs / / 1 / Enlarged femoral scales present (1) or absent (0) 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 No. of 4th toe lamellae 25 25 26 28 Sex f f m f TL 53.6 23.5 11.9/17.0 15.8/23.2 TW 4.1 4.1 4 5.1 FL 6.8 6.8 7.2 6.6 TBL 7.9 8.3 8.5 8.1 AG 18 18.7 20.2 18.3 HL 10.7 10.5 11 10.3 HW 6.2 6.2 7.1 6.4 HD 4 4.1 4.5 4.4 ED 2.2 2.5 2.4 2.3 EE 2.9 2.7 2.9 2.8 ES 5 4.8 5.5 5.1 EN 3.8 3.6 3.9 3.5 IO 2.2 2.6 2.9 2.4 EL 1 1 1.3 1.1 IN 0.8 1.1 1.2 1

Comparisons. Cnemaspis omari sp. nov. is one of five confirmed species in the siamensis group (Fig. 2). Within this group, it forms a monophyletic lineage with C. chanardi and its sister species C. roticanai. This lineage is diagnosed by the presence of a light colored prescapular crescent. Cnemaspis omari sp. nov. most closely resembles C. roticanai but differs from it in being smaller (max SVL 41.3 vs 47.0); having a lateral row of caudal tubercles; caudal tubercles that encircle the tail; and having a sequence divergence of 7.2% from C. roticanai (Table 4). It differs from C. chanardi in having four as opposed to 6–8 pore-bearing precloacal scales; and lacking a median row of enlarged subcaudal scales. From C. siamensis, C. omari is separated by having as opposed to lacking precloacal pores; having caudal tubercles that encircle the tail; lacking a row of enlarged median subcaudal scales; having a light colored pre-scapular crescent; and lacking dark, gular blotches. Form C. huaseesom, C. omari

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 57 93 sp. nov. differs in having keeled as opposed to smooth ventral scales; having four pore-bearing scales with round pores as opposed to 5–8 pore-bearing scales with elongate pores; having keeled as opposed to smooth subcaudal and subtibial scales; having caudal tubercles that encircle the tail; having a light-colored pre-scapular crescent; and lacking a yellow head, forelimbs, and tail in adult males. Etymology. We name this species in honor of the current Vice-Chancellor of Universiti Sains Malaysia, Penang, Professor Dato’ Omar Osman. This is a sign of appreciation for all the support and funding from the university and for accelerating the research of biodiversity and wildlife studies in Peninsular Malaysia for many years. Natural history. At the Phuphaphet Cave area in Satun, Thailand at 220 m in elevation, Grismer et al. (2010a) reported lizards being collected and observed during the day on the buttresses of trees and within tree holes between 1.5–2 m above the ground along a footpath in old, secondary forest. All the trees upon which the lizards were observed had holes into which the lizards would retreat upon provocation. Several rock outcrops were nearby but no lizards were observed on them. We have made similar observations on lizards from Perlis, Malaysia (Fig. 27), observing them at night on the trunks of large trees. LSUHC 9564 from the Phuphaphet Cave area was carrying two eggs indicating that the reproductive season extends through October.

FIGURE 27. Lowland dipterocarp forest of Cnemaspis omari sp. nov. at Perlis State Park, Perlis, Peninsular Malaysia. Photograph by LLG.

Cnemaspis roticanai Grismer & Chan, 2010 Roti Canai Rock Gecko Fig. 28

Holotype. ZRC 2.6860. Type locality: “743 m a.s.l. on Gunung Raya, Pulau Langkawi, Kedah, Peninsular Malaysia (06°22.114N, 99°49.270 E)” at 790 m in elevation. Diagnosis. Maximum SVL 47.0 mm; eight or nine supralabials; seven or eight infralabials; keeled ventral scales; 3–6 discontinuous, pore-bearing precloacal scales with round pores; 25–27 paravertebral tubercles; tubercles on flanks; tubercles in lateral caudal furrows; no ventrolateral caudal tubercles; caudal tubercles do not encircle tail; lateral caudal tubercle row absent; subcaudals keeled; median row of weakly enlarged subcaudal scales present; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 26–29 subdigital fourth toe lamellae; light colored prescapular crescent; gular and pectoral regions, abdomen, underside of hind limbs, and subcaudal region yellow in males (Tables 6,7).

58 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 94 FIGURE 28. Cnemaspis roticanai from Gunung Raya, Pulau Langkawi, Kedah, Peninsular Malaysia. Upper left: ventral coloration of adult male (lower) and adult female (upper) showing sexual dimorphism (LSUDPC 5192). Upper right: adult male (LSUDPC 5184) in light color pattern phase. Lower right: adult female (LSUDPC 5189) in dark color pattern phase. Lower left: microhabitat on Gunung Raya. Photographs by LLG.

Color pattern (Fig. 28). Males: dorsal ground color of head, body, limbs and tail pale-yellow; faint, brownish markings on top of head; single, diffuse, black, postorbital stripe extending to occiput and terminating at oblong, longitudinally oriented, yellow spot on nape followed by a pair of black, paravertebral spots on neck; rhomboidially shaped, brown, transverse markings between limb insertions extending onto tail as zig-zag-shaped, caudal bands; butterfly-shaped, pale-yellow interspaces between rhomboid markings; yellow, prescapular crescent followed by semi-transversely arranged, yellow bars on flanks separated by dark makings; forelimbs bearing yellowish blotches and scattered dark markings; hind limbs bearing yellowish blotches and dark markings resembling banding pattern; gular region yellowish orange; throat beige; abdomen, ventral surface of hind limbs, and subcaudal region yellow; ventral surface of forelimbs beige; all ventral scales bearing small, black stippling. Sexual dimorphism is marked in this species. Females: darker in overall coloration with much less yellow and a more contrasted dorsal pattern; dark markings on trunk appear as paravertebral blotches and in strong contrast to the pale-yellow interspaces; tail strongly banded; ventral surfaces yellowish throughout. Distribution. Cnemaspis roticanai is known only from Gunung Raya on Pulau Langkawi, Kedah, Peninsular Malaysia (Fig. 3). This species is expected to range more widely throughout the island. Natural history. Cnemaspis roticanai is a scansorial species occurring in open habitats with small, widely scattered rocks in hill dipterocarp forest above 400 m in elevation (Fig. 28). Grismer & Chan (2010) observed specimens on the underside of leaves, on tree trunks, and within cement drains. We hypothesize here that C. roticanai is a diurnal species being that the specimens we have observed at night were inactive and sleeping on tree trunks and the undersides of leaves. Others were found between rocks. Relationships. Cnemaspis roticanai is the sister species of C. omari sp. nov. (Fig. 2). Material examined. Malaysia: Kedah, Pulau Langkawi, Gunung Raya LSUHC 9453, ZRC 2.6860–62 (type series). Material examined since Grismer & Chan (2010): Kedah, Pulau Langkawi, Gunung Raya LSUHC 9430–31, 9439, 10802.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 59 95 Argus group. The argus group contains four species with a somewhat anomalous distribution pattern (Fig. 3). The basal species, Cnemaspis flavigaster Chan & Grismer occurs in central Peninsular Malaysia west of the Banjaran Titiwangsa mountains on the outskirts of Kuala Lumpur in Kepong, Ulu Gombak, and Batu Caves, Selangor (Chan & Grismer 2008) whereas the remaining three species, C. argus Dring and the sister species C. karsticola Grismer, Grismer, Wood & Chan and C. perhentianensis Grismer & Chan have restricted distributions in northeastern Peninsular Malaysia. Cnemaspis argus is known only from the mountainous region of Gunung Lawit (Dring 1979) and Gunung Tebu, Terengganu (Grismer et al. 2013c), C. karsticola is known from a single tower karst formation at Gunung Reng (Grismer et al. 2008b), and C. perhentianensis is endemic to two islands in the Perhentian Archipelago, Kelantan (Grismer & Chan 2008). The argus group is the sister lineage to the affinis group of central Peninsular Malaysia (Fig. 2) and is diagnosed by having a maximum SVL of 47.0–65.2 mm; 7–10 supralabials; 6–10 infralabials; 6–10 contiguous, pore-bearing, precloacal scales with round pores; randomly arranged dorsal tubercles extending onto the lower flanks; 17–32 paravertebral tubercles; no caudal tubercles in lateral furrows; a lateral row of caudal tubercles; no tubercles encircling the tail; no ventrolateral caudal tubercles; no median row of enlarged or keeled subcaudal scales; 1–4 postcloacal tubercles on each side of the base of the tail; no enlarged femoral or subtibial scales; subtibials keeled; and 27–35 lamellae beneath the fourth toe.

Cnemaspis flavigaster Chan & Grismer, 2008 Orange-bellied Rock Gecko Fig. 29

Cnemaspis affinis Boulenger 1912:39 Cnemaspis kumpoli Dring 1979:223

Holotype. HC 00282. Type locality: “the canopy walk trail at Forest Research Institute Malaysia (FRIM), state of Selangor, Peninsular Malaysia (3°14’23.04”N, 101°37’59.80” E)” at 120 m in elevation. Diagnosis. Maximum SVL 50.1 mm; nine or ten supralabials; 8–10 infralabials; smooth ventral scales; seven or eight contiguous pore-bearing precloacal scales with round pores; 21–24 paravertebral tubercles; tubercles on flanks; tubercles absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral caudal row of tubercles present; caudal tubercles do not encircle tail; subcaudals smooth with no enlarged, median scale row; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; enlarged submetatarsal scales on first toe; 29–34 subdigital fourth toe lamellae; wide, black, oblique stripe in shoulder region; distinct black and white bands on tail; pectoral region, abdomen, ventral surface of hind limbs, and subcaudal region orange in males (Tables 6,7). Color pattern (Fig. 29). Dorsal ground color brown to dark grey; sides of the head bearing yellow markings; three faint, postorbital stripes radiate from eye; three dark spots across the occiput anteriorly with another elliptical, medial spot posteriorly; paired, paravertebral spots on nape; dark, dorsolateral line extending from nape to forelimb insertion and onto brachium; paired, paravertebral, dorsal spots on body alternating with large, whitish blotches extending from shoulder to base of tail; another series of spots on flanks; flanks bear irregularly shaped, yellow blotches; dorsal surface of limbs bear irregularly shaped, dark and light markings; black and dull white bands nearly encircle tail; undersides of head and limbs mottled with fine, dark stippling; throat whitish; and chest, abdomen and underside of tail orange in males and beige in females. Coloration lightens considerably at night, highlighting the blotched dorsal pattern. Distribution. Cnemaspis flavigaster is known only from the Forest Research Institute of Malaysia, Batu Caves, Selangor (Chan & Grismer 2008), and is newly reported here from Ulu Gombak, Selangor (Fig. 3). Natural history. Cnemaspis flavigaster is a scansorial species restricted to granite rocks or karst (Flower 1899; Grismer & Chan 2008; Fig. 29) in lowland, old secondary forest and is only occasionally seen on tree trunks or fallen logs but never on leafy vegetation (Grismer 2011a). Grismer (2011a) noted that C. flavigaster is quite wary while abroad during the day on the shaded sides of rocks and deep within small alcoves. At night however, lizards venture farther out into open areas on the rocks and are more stationary. Grismer et al. (2010c) outlined a number of examples where sympatric species of Cnemaspis seem to be partitioning their habitat by having different activity periods, body sizes, substrate preferences or varying combinations of each. Inconsistent with

60 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 96 these observations was finding C. flavigaster at Ulu Gombak in syntopy on granite boulders with C. peninsularis sp. nov., Cnemaspis peninsularis sp. nov., however, is a habitat generalist and its presence on other substrates at Ulu Gombak may be enough to not preclude strong competition for resources. At Batu Caves, C. flavigaster has been observed at the entrance to Dark Cave. Relationships. Cnemaspis flavigaster is the basal species of the argus group (Fig. 2). Material examined. Malaysia: Selangor, Kepong, Forest Research Institute of Malaysia HC 00282, 00286; ZRC 2.6708–11 (type series). Material examined since Chan & Grismer (2008): Malaysia: Selangor, Kepong, Forest Research Institute of Malaysia LSUHC 6562, 8835–36; Ulu Gombak LSUHC 10380.

FIGURE 29. Cnemaspis flavigaster from FRIM, Kepong, Selangor, Peninsular Malaysia. Upper left: adult male (LSUDPC 4173) in the dark color pattern phase. Upper right: ventral view of adult male (LSUDPC 4181). Photographs by LLG. Middle left: adult female (LSUDPC 4601) in the dark color pattern phase. Photograph by CKO. Middle right: adult male (LSUDPC 4671) in the light color pattern phase. Lower left: subadult (LSUDPC 4178) in dark color pattern phase. Lower right; karst microhabitat at the Batu Caves, Selangor, Peninsular Malaysia. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 61 97 Cnemaspis argus Dring, 1979 Argus Rock Gecko Fig. 30

Holotype. BM 1974.4911. Type locality: “790 m on the east ridge of Gunung Lawit”, Terengganu, Peninsular Malaysia. Diagnosis. Maximum SVL 65.2 mm; eight or nine supralabials; eight or nine infralabials; keeled ventral scales; 6–10 pore-bearing precloacal scales; 26–32 paravertebral tubercles; tubercles not linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; no ventrolateral caudal tubercles, lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals keeled, no enlarged median scale row; 1–4 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 31–35 subdigital fourth toe lamellae; distinct black and white bands on tail (Tables 6,7). Color pattern (Fig. 30). Dorsal ground color greyish yellow; interorbital region green; yellow markings on head; paired, black, paravertebral, subelliptically shaped blotches extending from nape to tail and transforming into black bands; large patches of yellow tubercles and short transverse bars on flanks; limbs bearing faded, alternating, dark and light bands; non-regenerated tail bearing black and white bands; all ventral surfaces dull-white.

FIGURE 30. Left: microhabitat of Cnemaspis argus on Gunung Lawit, Terengganu, Peninsular Malaysia. Upper right: adult male (LSUDPC 6564) from Gunung Tebu, Terengganu, Peninsular Malaysia in the light color pattern phase. Photographs by LLG. Lower right: adult female (LSUDPC 6390) from Gunung Tebu in the dark color pattern phase. Photograph by ESHQ.

Distribution. Cnemaspis argus is known from Gunung Lawit (Dring 1979) and newly reported here from Gunung Tebu, Terengganu 10 km to the north along the same mountain range (Fig. 3). Natural history. Dring (1979) reported Cnemaspis argus to occur at 790 m in elevation in primary forest from Gunung Lawit and Grismer (2011a) considered it an upland endemic that would probably never be seen again because the trail up to Gunung Lawit had become overgrown and lost. However, we did find additional populations from the base of Gunung Lawit at 230 m in elevation and on Gunung Tebu from 40 m in elevation at Hutan Lipur Lata Belatan up to 750 m near the peak. All lizards were seen on large granite rocks within the forest (Fig. 30). These data indicate that C. argus is not an upland endemic but a microhabitat specialist restricted to granite rocks

62 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 98 wherever they may occur within its range. During the day, lizards remain wary and occur on the shady, vertical or inverted surfaces. They are in dark in overall coloration and difficult to approach. During the evening hours, lizards are much lighter in color and far less wary, tending to venture farther out onto the open areas of the boulders where they appear generally inactive. Relationships. Cnemaspis argus is most closely related to the sister species C. karsticola and C. perhentianensis (Fig. 2). Material examined. Malaysia: Terengganu, Gunung Lawit BM 1974.4910–11 (type series; photographs LSUDPC 2276–78), LSUHC 8304; Gunung Tebu LSUHC 10834–35, 10858–59.

FIGURE 31. Cnemaspis karsticola from Gunung Reng, Kelantan, Peninsular Malaysia. Upper left: adult female (LSUDPC 4424) in the dark color pattern phase. Upper right: adult male (LSUDPC 4421) in the dark color pattern phase. Lower; karst microhabitat at Gunung Reng. Photographs by LLG.

Cnemaspis karsticola Grismer, Grismer, Wood & Chan, 2008b Karst Rock Gecko Fig. 31

Holotype. ZRC 2.6765. Type locality: “Gunung Reng, Kelantan, Peninsular Malaysia (05°42.905 N, 101°44.726 E)” at 113 m in elevation.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 63 99 Diagnosis. Maximum SVL 48.1 mm; seven or eight supralabials; six or seven infralabials; ventral scales keeled; seven or eight contiguous, pore-bearing precloacal scales with round pores; 17–19 paravertebral tubercles; body tubercles not linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median scale row; two or three postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; 27–30 subdigital fourth toe lamellae; subcaudal region white (Tables 6,7). Color pattern (Fig. 31). Dorsal ground color yellowish brown; head and body overlain with faded, irregularly shaped, white markings; no dark, postorbital striping; three, dark, radiating, anteriorly projecting lines occur on occiput; paired, dark, paravertebral markings extend from nape to base of tail, similar markings on flanks; no transversely elongate, white markings on flanks; irregularly shaped, dark and light markings on limbs; diffuse, alternating, light markings on dorsal surface of tail; subcaudal region white, immaculate; regenerated portion of tail beige, immaculate. Distribution. Cnemaspis karsticola is known from only from Gunung Reng, Kelantan, Peninsular Malaysia (Grismer et al. 2008b). Natural history. Grismer et al. (2008b) noted that Cnemaspis karsticola is a lowland saxicolous species known only from the karst outcropping of Gunung Reng, a large isolated tower karst formation reaching 200 m in height situated along the east bank of the Pergau River at its junction with Batu Melintang, Kelantan. The base of the tower is undercut and numerous cracks and indentations accentuate and define its periphery (Fig. 31). Lizards were found during the day along the periphery of the karst formation and within cracks on the shaded surfaces of large, disconnected, karst boulders that have fragmented and fallen off the core. Lizards were not found deep within the larger cave system. Specimens were seen in cracks, on shaded overhangs, and on the cave walls no more than 2 m above the ground. No lizards were seen at night indicating this is a diurnal species. The color variation in the type series indicates this species is adept at substrate matching (Grismer et al. 2008b). Relationships. Cnemaspis karsticola is the sister species of C. perhentianensis Grismer & Chan (Fig. 2). Material examined. Malaysia: Kelantan, Gunung Reng ZRC 2.6763–65, LSUHC 9054. Additional material examined since Grismer et al. (2008b): Malaysia: Kelantan, Gunung Reng LSUHC 9053, 9055–56.

Cnemaspis perhentianensis Grismer & Chan, 2008 Perhentian Island Rock Gecko Fig. 32

Holotype. ZRC 2.6675. Type locality: “Pulau Perhentian Besar, Terengganu, West Malaysia (05°54.054 N, 102°44.726343 E)” at 40 m in elevation. Diagnosis. Maximum SVL 47.0 mm; 8–10 supralabials; seven or eight infralabials; ventral scales smooth to keeled; 6–8 contiguous, pore-bearing precloacal scales with round pores; 22–27 paravertebral tubercles; tubercles not linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present; caudal tubercles not encircling tail; all subcaudals keeled, no median row of enlarged subcaudals; three or four postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; 28–31 subdigital fourth toe lamellae; distinct black and white bands on tail (Tables 6,7). Color pattern (Fig. 32). Dorsal ground color grey to brown, overlain by irregularly shaped, white markings on top of head and snout; paired, white markings on occiput; no postorbital stripes; squarish, medial, white marking on neck; distinct, irregularly shaped, paravertebral, white markings on dorsum extending from shoulder region to base of tail and alternating with transversely elongate, distinct, white markings on flanks; three small, elongate, dark blotches at base of occiput; paired, dark, poorly defined, paravertebral blotches extend from nape to anterior portion of tail alternating with light, paravertebral markings; dark blotches on flanks alternating with white, transverse markings; black and dull white bands nearly encircle tail; irregularly shaped, dark and light markings on limbs; ventral surfaces of neck, body, and limbs beige, immaculate; gular region smudged with dark stippling.

64 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 100 FIGURE 32. Upper: adult male Cnemaspis perhentianensis (LSUDPC 5159) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia in the dark color pattern phase. Photographs by LLG. Lower: granite boulder microhabitat on Pulau Perhentian Besar. Photograph by Peter Carin.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 65 101 Distribution. Cnemaspis perhentianensis is known from only from Perhentian Besar and Perhentian Kecil islands (Grismer & Chan 2008; Grismer et al. 2011a) of the Perhentian Archipelago, Terengganu off the northeast coast of Peninsular Malaysia (Fig. 3). Natural history. The Perhentian Archipelago is composed of 11 relatively small islands lying 21 km off the east coast of the state of Terengganu (Fig. 3). The largest of these islands, Pulau Perhentian Besar (ca. 857 hectares) is a rugged, hilly island reaching 249 m in elevation. The majority of the island is covered in primary, lowland dipterocarp forest and its granite bedrock is the source of extensive boulder outcroppings that add a significant degree of habitat and microhabitat complexity to the island’s ecosystem, which in turn, supports various saxicolous species (Grismer et al. 2011a). Grismer & Chan (2008) noted that Cnemaspis perhentianensis is restricted solely to the granite outcroppings (Fig. 32). Specimens collected or observed during the day were found on the shaded surfaces of both large and small rocks and would retreat into cracks at the slightest provocation. Upon retreat, lizards would roll their tail over their back and wag the tip from side to side. At night, lizards could be found on all surfaces of the rocks at greater distances from their crevice microhabitats and appreaed inactive. No lizards were seen on tree trunks or other types of vegetation. Grismer & Chan (2008) reported that the activity of C. perhentianensis may be closely tied to precipitation, noting that on multiple trips, specimens were only observed following periods of rain. Relationships. Cnemaspis perhentianensis is the sister species of C. karsticola (Fig. 2). Material examined. Malaysia: Terengganu, Pulau Perhentian Besar ZRC 2.6675–79 (type series). Additional material examined since Grismer & Chan (2008): Malaysia: Terengganu, Pulau Perhentian Besar LSUHC 8673, 8675–76, 8697–700, 9060, 9412. Affinis group. The affinis group contains 13 species that collectively range throughout central Peninsular Malaysia and is comprised of three basal lineages: Cnemaspis pseudomcguirei from northwestern Peninsular Malaysia; the sister species C. affinis and C. harimau from northwestern Peninsular Malaysia and their sister lineage that contains the remaining 10 species (Figs. 2,3). The latter lineage, for the most part, is a polytomy. However, the sister species relationship between C. mcguirei and C. grismeri associated with the Banjaran Bintang Mountains and the relationships between the diminutive, lowland karst-dwellers C. hangus sp. nov., C. selamatkanmerapoh, and C. bayuensis east of the Banjaran Titiwangsa Mountains and the small, upland granite- dweller C. stongensis sp. nov. in the northern Banjaran Titiwangsa Mountains are strongly supported (Fig. 2). The sister species relationship between C. mcguirei and C. grismeri is strongly supported in the molecular analysis and further supported in the morphological analysis by them having a pair of ocelli in the shoulder region which we consider to be a derived condition in that it occurs in no other species of Cnemaspis. The molecular analysis indicates that Cnemaspis flavolineata from the type locality at the Gap below Fraser’s Hill, Pahang is not conspecific with C. flavolineata from Cameron Highlands (18.0% sequence divergence; Table 4), 76 km to the north and the two species may not even be closely related (Fig. 2). We also found significant differences in tuberculation separating these two populations. Smith (1922) was the first to report this species from Fraser’s Hill which he considered conspecific with C. kendallii (Gray) but later (Smith 1930) and without comment, referred to it as C. affinis. Although Nicholls (1949) was the first to recognize this population as a distinct species, his brief description did not provide a single diagnostic character separating it from either C. affinis or C. kendallii (quite frankly, he got lucky). As such we provide a redescription of this species below based on the holotype and a second specimen recently collected from the type locality and describe the Cameron Highlands population as a new species (C. temiah sp. nov.). The affinis group is diagnosed as having a maximum SVL of 36.5–65.0 mm; 7–13 supralabials; 7–11 infralabials; keeled ventral scales; 0–10 pore-bearing precloacal scales with round pores; 18–34 paravertebral tubercles; tubercles on flanks; lateral row of caudal tubercles present; subcaudals keeled; no median row of enlarged subcaudal scales; 1–5 postcloacal tubercles on each side of base of tail; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; and 21–35 lamellae beneath the fourth toe.

Cnemaspis affinis (Stoliczka, 1870) Penang Island Rock Gecko Fig. 33

Gymnodactylus affinis Boulenger 1885:42 Gonatodes penangensis Flower 1896:863

66 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 102 FIGURE 33. Cnemaspis affinis from Penang Hill, Pulau Pinang, Penang, Peninsular Malaysia. Upper left: adult male (LSUDPC 9078) in the dark color pattern phase. Lower left: adult female (LSUDPC 9079) in the light color pattern phase. Upper right: microhabitat of C. affinis on Penang Hill. Lower right: ventral coloration of male (left) and female (right) C. affinis. Photographs by LLG.

Holotype. ZSI 5964. Type locality: “Penang Hills”, Penang Island, Penang, Peninsular Malaysia at approximately 800 m in elevation. Diagnosis. Maximum SVL 50.8 mm; 9–13 supralabials; 8–10 infralabials; ventral scales keeled; five or six discontinuous, pore-bearing precloacal scales with round pores; 20–28 paravertebral tubercles; dorsal tubercles not linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; ventrolateral caudal tubercles absent; lateral caudal row present; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median scale row; two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; 28 or 29 subdigital fourth toe lamellae; gular region yellow in males; single ocellus in shoulder region in males, yellow post-scapular band in males; and transverse yellow bars on flanks (Tables 6,7). Color pattern in life (Fig. 33). Adult males: dorsal ground color grey to brown; paired white markings on occiput; dark pre- and postorbital stripes are present with latter extending onto nape; medial, white marking on nape followed by distinct, large, black shoulder patches in males usually enclosing a white to yellow ocellus anteriorly and edged posteriorly by yellow spot; irregularly shaped, paravertebral, white markings on dorsum

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 67 103 extending to base of tail and confluent with transversely elongate, distinct, yellow markings on flank. Adult females: slightly less boldly marked; tend to lack postscapular band, more yellowish overall. Distribution. Cnemaspis affinis is endemic to Penang Island, Penang, Peninsular Malaysia (Grismer et al. 2008b; Fig. 3). Natural history. Grismer et al. (2008b) indicated that Cnemaspis affinis is found only in the upland regions of Penang Island in the vicinity of 720 m where it occurs on large granite boulders in disturbed forest (Fig. 33). During the day, lizards are active on the shaded surfaces of the boulders and quite wary but at night, venture farther out onto the boulder’s surface (Flower 1896). Flower (1896) reported finding a specimen beneath the bark of a large tree. We suspect, however, this was not C. affinis but the similarly appearing C. shahruli, which is a habitat generalist and known to occur on vegetation (Grismer et al. 2010c; Grismer 2011a). Gravid females carrying two eggs have been observed during July. Relationships. Cnemaspis affinis is the sister species of C. harimau (Fig. 2). Material examined. Malaysia: Penang, Penang Island LSUHC 6758–59, 6773–74, 6787–88, ZRC 2.1098, 2.5203, 2.4858, 2.6017–18, ZSI 5964 (holotype). Additional material examined since Grismer & Chan (2008): Malaysia: Penang, Penang Island LSUHC 10347.

Cnemaspis harimau Chan, Grismer, Shahrul, Quah, Muin, Savage, Grismer, Norhayati, Remegio & Greer, 2010b Tiger Rock Gecko Fig. 34

Holotype. ZRC 2.6894. Type locality: “Sungai Badak (=Badak river), Gunung Jerai, Kedah, Peninsular Malaysia (N 05°48.59’, E 100°23.53’)” at 600 m in elevation.

FIGURE 34. Cnemaspis harimau from Sungai Badak, Gunung Jerai, Kedah, Peninsular Malaysia. Upper left: adult male (LSUDPC 6381) in the dark color pattern phase. Upper right: adult female (LSUDPC 5334) in the light color pattern phase. Lower left: adult female (LSUDPC 6382) in the light color pattern phase. Photographs by LLG. Middle right: juvenile male (LSUDPC 6520) in light color pattern phase. Photograph by ESHQ. Lower right: ventral view of adult male (LSUDPC 5337). Photograph by LLG.

68 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 104 FIGURE 35. Microhabitat of Cnemaspis harimau at Sungai Badak, Gunung Jerai, Kedah, Peninsular Malaysia. Photograph by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 69 105 Diagnosis. Maximum SVL 40.7 mm; nine or 10 supralabials; nine or 10 infralabials; ventral scales keeled; four discontinuous, pore-bearing precloacal scales with round pores; 18–20 paravertebral tubercles; tubercles not linearly arranged, present on flanks; tubercles in lateral caudal furrows; ventrolateral caudal tubercles absent anteriorly; lateral row of caudal tubercles present; caudal tubercles encircling tail; all subcaudals keeled, no enlarged median scale row; two or three postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 25–30 subdigital fourth toe lamellae; gular region and throat yellow in males; single ocellus in shoulder region in males; yellow postscapular band variable; transverse yellow bars on flanks (Tables 6,7). Color pattern in life (Fig. 34). Ground color of dorsum grayish-brown; top of head yellowish with an indistinct, gray speckling pattern on occiput; faint, preorbital stripe; three black lines radiate from anterior region of nape onto base of occiput; light, gray, irregularly shaped, paravertebral blotches extend from nape to base of tail; broken, dark, paravertebral, longitudinal streaks begin with paired streaks in prescapular region and alternate with paravertebral blotches posteriorly, terminating at base of tail; black shoulder patch bordered anteriorly by short, yellow band enclosing a whitish ocellus and bordered posteriorly by a longer, yellow band; shoulder patch absent in females; a series of yellow bands on flanks decreasing in length posteriorly with last band terminating just anterior to hind limb insertion; dorsal surfaces of limbs speckled with dark and yellowish markings; tail distinctly marked with diffuse, dark-gray and whitish bands; gular region and throat yellow-orange in males; ventral surfaces of limbs, remainder of body, and base of tail light gray; and subcaudal region darkly stippled. Distribution. Cnemaspis harimau is endemic to Gunung Jerai, Kedah in northwestern Peninsular Malaysia (Chan et al. 2010b; Fig. 3). Natural history. Gunung Jerai is an isolated mountain reaching 1217 m in elevation on the northwest shore of Peninsular Malaysia. Chan et al. (2010b) reported finding lizards on granite rocks and at the base of trees at 600 m in elevation during the day and night in hill forest vegetation (Fig. 35). Lizards appear to be far more active during the day and are much more sedentary at night. When frightened, lizards will seek refuge in the porus matrices formed by the dry expansive soil at the base of the rocks (Grismer 2011a). We have observed lizards on isolated rocks along the edge of the main road leading to the summit of Gunung Jerai beginning at approximately 400 m in elevation. Relationships. Cnemaspis harimau is the sister species of C. affinis (Fig. 2). Material examined. Malaysia: Kedah, Gunung Jeari LSUHC 9665, 9667, 9669, ZRC 2.6894–97 (type series). Additional material examined since Chan et al. (2010): Malaysia: Kedah, Gunung Jerai LSUHC 8229, 8232–33, 9666, 9668.

Cnemaspis pseudomcguirei Grismer, Norhayati, Chan, Belabut, Muin, Wood & Grismer, 2009 False-McGuire Rock Gecko Fig. 36

Holotype. ZRC 2.6777. Type locality: “Bukit Larut, Perak, Peninsular Malaysia along the road from the Gunung Hijau Rest House to the Telekom Tower (04°51.715 N, 100°47.993 E)” at 1351 m in elevation. Diagnosis. Maximum SVL 42.5 mm; nine or 10 supralabials; 8–10 infralabials; ventral scales keeled; 1–5 continuous, pore-bearing precloacal scales with round pores; 23–32 paravertebral tubercles; body tubercles randomly arranged, present on flanks; tubercles present in lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present anteriorly; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median scale row; two or three postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; usually no enlarged submetatarsal scales on first toe; 23–26 subdigital fourth toe lamellae; and two ocelli in the shoulder region in males; vertebral stripe variable (Tables 6,7). Color pattern in life (Fig. 36). Dorsal ground color of head, body, limbs and tail golden brown; top of head bearing small, black and yellow markings; thin, black, postorbital stripe; thin, yellow, postorbital stripes unite on nape and may continue posteriorly to form a wide, vertebral stripe that fades in pelvic region; vertebral stripe variably present, often replaced by white, paravertebral markings edged anteriorly in black; dark markings on nape anterior to yellow stripe; black shoulder patch encloses a larger, upper, posterior white ocellus and smaller, lower, anterior ocellus; single, light, postscapular spot posterior to black shoulder patches; faint, yellow, transverse

70 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 106 markings on flanks; faint, dark and dull-yellow reticulum on limbs; tail faintly marked with light-brown and dull- yellow bands; all ventral surfaces beige with weak stippling. The color pattern lightens considerably at night and many lizards appear nearly unicolor light-yellow to white.

FIGURE 36. Cnemaspis pseudomcguirei from Bukit Larut, Perak, Peninsular Malaysia. Upper left: adult male (LSUDPC 4723) in the dark color pattern phase. Upper right: adult female (LSUDPC 4756) in light color pattern phase sleeping on a leaf at night. Lower left: gravid female (LSUDPC 4731) in the dark color pattern phase. Lower right: hatchling (LSUDPC 6397). Photographs by LLG.

Distribution. Cnemaspis pseudomcguirei is known from Bukit Larut and Gunung Inas in the Banjaran Bintang Mountains, Perak, in northwestern Peninsular Malaysia (Grismer et al. 2009; Laidlaw 1901; Fig. 3). Natural history. Grismer (2011a) noted that Cnemaspis pseudomcguirei is an upland species found in hill dipterocarp forests from at least 1,000–1,300 meters in elevation (Fig. 37). During the day, lizards are found beneath small stones and logs on the forest floor or within rotten logs. Unlike the sympatric species C. mcguirei, C. pseudomcguirei does not occur on large granite boulders and lizards are commonly found great distances (>1 km) from boulder outcroppings. At Gunung Inas, Laidlaw (1901) reported lizards being numerous “amongst boulders on the course of a small stream” which we believe he confused with the sympatric and similarly appearing C. mcguirei. During the night, C. pseudomcguirei can be found exposed sleeping on the surfaces of leaves 0.5–1.5 m above the ground. Females carrying two eggs have been found during March and October suggesting this species breeds through the rainy season. Relationships. Cnemaspis pseudomcguirei is part of the basal tritomy of the affinis group (Fig. 2). Material examined. Malaysia: Perak; Bukit Larut ZRC 2.6777–82 (type series). Material examined since Grismer et al. (2009): Malaysia: Perak, Bukit Larut LSUHC 9074, 10640, 10645.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 71 107 FIGURE 37. Microhabitat of Cnemaspis pseudomcguirei at Bukit Larut, Perak, Peninsular Malaysia. Photograph by LLG.

Cnemaspis shahruli Grismer, Chan, Quah, Mohd, Savage, Grismer,Norhayati, Greer & Remegio, 2010c

Shahrul’s Rock Gecko Fig. 38

Holotype. ZRC 2.6898. Type locality: “Telok Bahang Recreational Forest Reserve, Penang (05°27.233’N, 100°12.324’E)” at 67 m in elevation. Diagnosis. Maximum SVL 36.5 mm; 10 or 11 supralabials; 8–10 infralabials; ventral scales keeled; no precloacal pores; 19–23 paravertebral tubercles; body tubercles randomly arranged, present on flanks; tubercles within lateral caudal furrows; ventrolateral caudal tubercles absent anteriorly; lateral row of caudal tubercles present anteriorly; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median row; 1–3 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; usually no enlarged submetatarsal scales on first toe; 21–30 subdigital fourth toe lamellae; light-colored vertebral stripe variably present; gular region, throat and pectoral region yellow in males; dark, central, elongate marking occurs in mental region; a single ocellus in the shoulder region in males; white, dorsal, caudal tubercles; distinct black and white caudal bands variably present (Tables 6,7). Color pattern in life (Fig. 38). Dorsal ground color of head, body, limbs and tail pale brown; top of head with dark and light diffuse markings; dark, postorbital blotches; when present, a wide, light colored, vertebral stripe extends from occiput to base of tail; offset, opposing, paravertebral, dark markings extend from nape to base of tail; large, black, square, shoulder patches enclose a whitish to yellow ocellus in adult males, absent in females and juveniles; dark shoulder patches become greatly reduced in the nighttime color phase; large, light-colored markings occur on flanks and tend to form transverse bands; forelimbs and hind limbs mottled; gular region, throat, and

72 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 108 anterior pectoral region yellow; gular region often faintly mottled; a dark, central, elongate marking occurs in mental region; ventral surfaces of limbs and body beige with small, black stipples in each scale; subcaudal region darker; tail faintly banded; regenerated tail mottled.

FIGURE 38. Cnemaspis shahruli from Peninsular Malaysia. Upper left: microhabitat of C. shahruli on Pulau Pangkor, Perak. Upper right: adult male (LSUDPC 5320) from Pulau Pinang, Penang in the dark color pattern phase. Photographs by LLG. Middle right: adult female (LSUDPC 6304) from Pulau Pangkor in the dark color pattern phase. Photograph by J. van Rooijen. Lower right: juvenile (LSUDPC 5880) from Bukit Mertajam, Penang in the light color pattern phase. Photograph by ESHQ. Lower left: ventral coloration of adult male (LSUDPC 5317) from Pulau Pinang.

Distribution. In northwestern Peninsular Malaysia, Cnemaspis shahruli is known from Penang Island and the adjacent island of Jerejak, Penang and the mainland localities of Sungai Sedim, Kedah and Bukit Mertajam, Penang. One hundred thirteen km farther south, C. shahruli is known from Pulau Pangkor, Perak, (Grismer 2011a; Grismer et al. 2010c; Fig. 3) and it is expected to occur in the intervening mainland areas between Pulau Pangkor and Bukit Mertajam. Natural history. Cnemaspis shahruli is generally a nocturnal species that is only occasionally seen during the day. It is also a habitat generalist that occurs in both riparian and non-riparian microhabitats from near sea level to approximately 500 m in elevation and can be found on both rocks and vegetation (Grismer et al. 2010c; Fig. 38). In lowland areas on Penang Island, C. shahruli occurs in riparian areas where it is usually found on granite rocks and boulders that may or may not be covered in lichen or vegetation. Lizards are occasionally found on adjacent tree trunks provided they are near large rocks. On Pulau Jerejak, C. shahruli has been found in disturbed areas on rocks and tree trunks (Grismer et al. 2010c). Lizards are occasionally found beneath logs and at the base of trees. The only specimen known from Pulau Pangkor, Perak was collected during the day along a forest trail while it was clinging upside down to the underside of a leaf approximately 0.5 m above the ground.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 73 109 Relationships. Cnemaspis shahruli may be the sister species to C. narathiwatensis (Fig. 2) although this relation is not statistically supported. Material examined. Malaysia: Penang; Pulau Pinang ZRC 2.6898–902; Pulau Jerejak ZRC 2.6903; Sungai Sedim ZRC 2.6904. Perak: Pulau Pangkor ZRC 2.6905 (type series). Additional specimens examined since Grismer et al. (2010c): Malaysia: Penang; Pulau Pinang LSUHC 8875, 8879, 9572, 9575, 9595, 9862; Bukit Mertajam LSUHC 10375.

FIGURE 39. Adult male Cnemaspis mcguirei (LSUDPC 5173) from Bukit Larut, Perak, Peninsular Malaysia in the light color pattern phase. Photograph by LLG.

Cnemaspis mcguirei Grismer, Grismer, Wood & Chan, 2008b McGuire’s Rock Gecko Figs. 39, 40

Gonatodes affinis Laidlaw 1901:304; Boulenger 1903:148, 1912:38; Smith 1930:16 Gonatodes kendalli Boulenger 1912:38 Cnemaspis kendallii Das & Bauer 1998:13 (in part)

Holotype. ZRC 2.6765. Type locality: “Bukit Larut, Perak, Peninsular Malaysia (04°51.715 N, 100°47.993)” at 1351 m in elevation. Diagnosis. Maximum SVL 65.0 mm; 7–10 supralabials; 7–9 infralabials; ventral scales keeled; 5–10 usually discontinuous, pore-bearing precloacal scales with round pores; 26–32 paravertebral tubercles; body tubercles not linearly arranged, present on flanks; tubercles present in lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median subcaudal scale row; 2–5 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 27–35 subdigital fourth toe lamellae; two ocelli in the shoulder region in males; wide, white to yellow postscapular band; yellow bars on flanks; distinct black and white caudal bands posteriorly (Tables 6,7).

74 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 110 FIGURE 40. Cnemaspis mcguirei (LSUDPC 5173) from Bukit Larut, Perak, Peninsular Malaysia. Upper left: adult male (LSUDPC 6303) in the dark color pattern phase. Photograph by CKO. Upper right: adult male (LSUDPC 5175) in the dark color pattern phase illustrating the dorsal ocelli. Lower right: adult female (LSUDPC 6409) in the dark color pattern phase in the typical diurnal position of clinging upside down to the underside of an overhang on a granite rock. Lower left: granite boulder microhabitat at Bukit Larut. Photographs by LLG.

Color pattern (Figs. 39, 40). Dorsal ground color grey to brown; head and body overlain with irregularly shaped, dark blotches; light markings on top of head; dark, postorbital stripe extends onto nape and contacts dark, anteriorly projecting, median stripe; medial, white marking occurs on nape followed by distinct, large, black, shoulder patches in males enclosing one or two (usually two) yellow ocelli; shoulder patches edged posteriorly by wide, postscapular band that is yellow laterally and white dorsally; irregularly shaped, paravertebral, white markings occur on body and extend to base of tail; transversely elongate, distinct, yellow markings occur on flanks; diffuse, brown and dull white bands encircle tail; irregularly shaped, dark and light markings occur on limbs; ventral surfaces of head, body, and limbs dull beige, immaculate and darkened laterally; females less boldly marked than males. Coloration lightens considerably at night. Distribution. Grismer et al. (2008b) noted that Cnemaspis mcguirei ranges through Banjaran Bintang Mountains (Fig. 3) beginning at an isolated population from Gunung Bubu, Perak in the south, northward through the continuous central section of the Banjaran Bintang Mountains from Bukit Larut to Gunung Inas (Laidlaw 1901; Boulenger 1912) in the north (Fig. 3). From here the Banjaran Bintang Mountains continue northward and merge with the more extensive Banjaran Titiwangsa Mountains and these continue farther north as a series of smaller, parallel, north to south tending, somewhat isolated ranges, terminating just south of Pattani, Pattani Province, Thailand. At Namtok Sai Khao on the northeast perimeter of this range, Boulenger (1903) provided descriptions of four specimens that match the description of C. mcguirei and Grismer et al. (2008b), based on the examination of photographs (LSUDPC 4581–82) of these specimens (BM 1903.4.15.11–14), considered them to represent the northern extent of C. mcguirei. We follow that taxonomy here but note that this presents a 200 km hiatus from its northernmost distribution at Gunung Inas and that a firsthand examination of these specimens and a molecular analysis of this population may indicate they are different species.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 75 111 FIGURE 41. Cnemaspis grismeri from Gua Asar, Bukit Kepala Gajah, Lenggong, Perak, Malaysia. Upper: adult female (LSUDPC 6693) in the dark color pattern phase. Lower: adult male (LSUDPC 6696) in the dark color pattern phase. Photographs by ESHQ.

76 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 112 FIGURE 42. Upper: juvenile Cnemaspis grismeri (LSUDPC 6695) from Gua Asar, Bukit Kepala Gajah limestone massif, Lenggong, Perak, Malaysia in light color pattern phase. Lower: karst microhabitat of at Gua Asar. Photographs by ESHQ.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 77 113 Natural history. Grismer (2011a) noted that Cnemaspis mcguirei occurs in rocky, hilly terrain from approximately 900–1,300 m in elevation in lowland and hill dipterocarp and lower montane forests and occurs almost exclusively on granite rocks (Fig. 40) devoid of moss or on the areas of rock where moss is not growing. Only occasionally are lizards found on logs or the mossy parts of rocks. Lizards are commonly seen during the day in heavily shaded areas at the edges of their retreats or clinging upside down to the undersides of large boulders (Fig. 40). In such microhabitats, lizards are cryptic and often noticed only by the yellow ocelli that appear as eye- spots. Lizards are wary and quickly move to deeper cover at the slightest provocation, often waving their tails from side to side over their backs as they retreat. At night, C. mcguirei takes on a much lighter coloration and ventures further out onto the surface of the rocks (Fig. 39) and is easily approached. Lizards have been observed abroad at night during heavy rains resting in dry, open patches on rocks sheltered from water. Several termites were found in the gut of one lizard and females carrying two eggs were observed during March and October suggesting C. mcguirei breeds throughout the year. Relationships. Cnemaspis mcguirei is the sister species of Cnemaspis grismeri (Fig. 2). Material examined. Malaysia: Perak, Bukit Larut ZRC 2.6765–69 (type series). Material examined since Grismer et al. (2008b): Malaysia: Perak, Bukit Larut ZRC 2.5663–64, LSUHC 8855, 8858, 9140, 9028–33, 9209, 9845; Gunung Bubu LSUHC 11013.

Cnemaspis grismeri Wood, Quah, Anuar & Muin, 2013 Grismer’s Rock Gecko Figs. 41,42

Cnemaspis mcguirei Grismer 2011a:349.

Holotype. LSUHC 10996. Type locality: “Gua Asar, Bukit Kepala Gajah limestone massif, Lenggong, Perak, Malaysia (5°07.53’N, 100°58.82’E)” at 78 m in elevation. Diagnosis. Maximum SVL 50.6 mm; eight supralabials and infralabials; ventral scales keeled; 8–10 discontinuous, pore-bearing precloacal scales with round pores; 27–32 paravertebral tubercles; body tubercles not linearly arranged, present on flanks; tubercles present in lateral caudal furrows; ventrolateral caudal tubercles present anteriorly; lateral row of caudal tubercles present; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median scale row; two or three postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; 25–31 subdigital fourth toe lamellae; two ocelli in the shoulder region in males; wide, white to yellow postscapular band; yellow bars on flanks; nearly immaculate white bands on tail (Tables 6,7). Color pattern (Figs. 41,42). Dorsal ground color grey to brown; head and body overlain with irregularly shaped, small, dark and yellowish flecks giving an overall mossy appearance; cream to yellowish markings on top of head; thin, dark, postorbital stripe extending onto nape; paired, elongate, medial, yellowish markings on nape followed by small, indistinct, black shoulder patches enclosing two yellow ocelli, ocellus dorsal to forelimb insertion distinct and another anterior to forelimb insertion weak; shoulder patch edged posteriorly by wide, offset, postscapular band that is yellow laterally and white medially; irregularly shaped, offset, paravertebral, yellowish markings on dorsum extend to base of tail; distinct, transversely elongate, yellow bars on flanks; diffuse, brown and white bands encircle tail posteriorly, bands yellowish anteriorly; posterior portion of tail regenerated and uniform dark brown; irregularly shaped yellowish to dull white markings on limbs; dark and light diffuse bands encircling digits; ventral surfaces of head, body, and limbs dull beige, immaculate, darkening laterally; subcaudal region suffused with pigment, not immaculate. Distribution. Cnemaspis grismeri is known only from the type locality at Gua Asar, Bukit Kepala Gajah limestone massif, Lenggong, Perak, Malaysia (Wood et al. 2013; Fig. 3). Natural history. Cnemaspis grismeri is a lowland species found on the inner walls of limestone caves, cave entrances, karst walls outside of caves, and outcroppings in the lowland karst forest surrounding limestone massifs at Gua Asar and Gua Kijang (Fig. 42). Wood et al. (2013) reported that C. grismeri is diurnal but only observed lizards moving about in cracks and on shadowed surfaces of karst boulders. Lizards are wary and quickly enter into retreats when approached. Their behavior is similar to that of its upland closest relative C. mcguirei that resides on granite boulders at Bukit Larut, Perak (Grismer 2011a). At night, the color of C. grismeri lightens considerably and

78 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 114 FIGURE 43. Upper: adult male Cnemaspis flavolineata (LSUHC 8079) from the Gap at Fraser’s Hill, Pahang, Peninsular Malaysia. Lower: microhabitat at Fraser’s Hill. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 79 115 lizards have been observed sleeping on leaves in low vegetation near the karst walls and on vines and the aerial roots of fig trees next to the limestone walls (Wood et al. 2013). Gravid females bearing two eggs have been found during July. Relationships. Cnemaspis grismeri is the sister species of C. mcguirei (Fig. 2) and both are closely associated with the Banjaran Bintang Mountains (Fig. 1), C. grismeri being the lowland form and C. mcguirei the upland species. A parallel phylogeographic pattern occurs in the upland Cyrtodactylus bintangtinggi Grismer, Wood, Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer & Pauwels and C. bintangrendah Grismer, Wood, Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer & Pauwels (Grismer et al. 2012, 2014) from the same localities. Material examined. Malaysia: Perak, Lenggong LSUHC 9969–73, 10941–44, 10996 (type series).

Cnemaspis flavolineata (Nicholls, 1949) Fraser’s Hill Rock Gecko Fig. 43

Gonatodes kendallii Smith 1922:268 Gonatodes affinis Smith 1925:23; 1930:16 Gonatodes flavolineatus Nicholls 1949:47 Cnemaspis flavolineatus Manthey & Grossmann 1997:211; Leong & Lim, 2003:9

Holotype. ZRC 2.6777. Type locality: “the Gap below Fraser’s Hill, on the Pahang-Selangor boundary”, Peninsular Malaysia at approximately 800 m in elevation. Diagnosis. Maximum SVL 39.2 mm; nine supralabials; nine infralabials; ventral scales keeled; five or six contiguous, pore-bearing precloacal scales with round pores; 23 paravertebral tubercles; body tubercles linearly arranged, absent on flanks; tubercles present in lateral caudal furrows; no ventrolateral caudal tubercles; lateral caudal row present anteriorly; caudal tubercles encircle tail anteriorly; subcaudals keeled, no enlarged median scale row; two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales; and 23 subdigital fourth toe lamellae; large, black, round spots on nape and anterior portion of body; light vertebral stripe variably present (Tables 6,7). Redescription of species. The redescription of this species is based on a specimen (LSUHC 8079) collected from the Gap below Fraser’s Hill, Pahang and the holotype (ZRC 2.6777). Maximum SVL 38.6–39.2 mm; head oblong in dorsal profile, moderate in size, somewhat narrow, flattened, distinct from neck; snout short, flat in lateral profile; postnasal region constricted medially, flat; scales of rostrum keeled, raised, larger than similarly shaped scales on occiput; weak, supraorbital ridges and frontorostral sulcus; canthus rostralis smoothly rounded; eye large; extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral slightly concave, divided dorsally by longitudinal groove; rostral bordered posteriorly by supranasals and two smaller scales and laterally by first supralabials; 9R,L raised supralabials of similar size; 9R,L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented posterolaterally, bordered posteriorly by small, granular, postnasal scales; mental large, triangular, bordered posteriorly by three postmentals, outer two largest; gular scales raised, keeled; throat scales larger, raised, keeled. Body slender; small, keeled, dorsal scales equal in size throughout body, intermixed with much larger, multicarinate tubercles more or less linearly arranged; tubercles extend from occiput to base of tail; no tubercles on flanks; 23 paravertebral tubercles; pectoral scales raised, keeled, not elongate; abdominal scales slightly larger than dorsals, flat, keeled; five or six precloacal pores; forelimbs moderately long, slender; dorsal scales of brachium raised, keeled; dorsal scales of forearm keeled, raised; ventral scales of brachium keeled, raised, juxtaposed; ventral scales of forearm smooth, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate

80 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 116 with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; toes increase in length from first to fourth with fourth being slightly longer than fifth; 23 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales raised anteriorly, weakly keeled, juxtaposed; middorsal and lateral caudal furrows; no row of enlarged, median subcaudal scales; subcaudal scales keeled; caudal tubercles encircle tail anteriorly; caudal tubercles present in lateral furrow; two enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail. Color pattern in life based on LSUHC 8079 (Fig. 43). Dorsal ground color brown to olive-brown; small, dark and light markings on top of head; faint, dark lines radiate out from eyes; elongate, paired, dark, paravertebral markings edging a wide, cream-colored vertebral stripe extending from nape to base of tail; flanks slightly lighter in color; large, white, dorsal tubercles in vertebral region extending from nape onto the tail; limbs mottled with dark and light, irregularly shaped blotches; no black shoulder patches or postscapular band; faint, dark and light brownish bands encircle tail; ventral surfaces of head, body, and limbs dull beige, immaculate; subcaudal region darker. Distribution. Cnemaspis flavolineata is known only from the type of the Gap below Fraser’s Hill, Pahang (Nicholls 1949; Fig. 3). Natural history. This species is known from only two specimens. No collection data were provided with the holotype but LSUHC 8079 was taken on 26 August 2006 at night while sleeping on the underside of a horizontal branch at the edge of a slow moving, marshy stream edged by large granite boulders in hill dipterocarp forest at 900 m in elevation (Fig. 43). Remarks. Although Cnemaspis flavolineata and C. temiah sp. nov. are very similar in morphology and color pattern, the molecular analysis indicates that they are not the same species or each others closest relatives (Fig. 2). Cnemaspis flavolineata can be separated from C. temiah sp. nov. by having a distinct series of large, white, transversely arranged, vertebral tubercles extending from the nape onto the tail which do not occur in C. temiah sp. nov. Additionally, the anteriormost caudal tubercles in C. flavolineata encircle the tail whereas no caudal tubercles encircle the tail in C. temiah sp. nov. Material examined. Malaysia: Pahang; the Gap below Fraser’s Hill ZRC 2.6777 (holotype) and LSUHC 8079.

Cnemaspis temiah sp. nov Temiah Rock Gecko Fig. 44

Cnemaspis affinis Grandison 1972:80; Dring 1979:221 Cnemaspis flavolineata Manthey & Grossmann 1997:211 (in part); Chan-ard et al 1999:104, Grismer 2008:30; Grismer et al. 2008c:9 (in part); Grismer 2011a:317 Cnemaspis flavolineatus Lim et al. 2002:51

Holotype. Adult female LSUHC 9110 collected on 11 November 2008 by L. Lee Grismer, Norhayati Ahmad, and Chan K. Onn on trail 11, Tanah Rata, Cameron Highlands, Pahang, Peninsular Malaysia (03°09.01 N, 106°14.03 E) at approximately 1600 m in elevation. Paratypes. All paratypes are from the same locality as the holotype. LSUHC 9159–60 have the same collectors and collection dates. LSUHC 9739 was collected by Chan K. Onn on 22 March 2010; LSUHC 9816–18 were collected by L. Lee Grismer, Chan K. Onn, and R. Gregory on 27 August 2010. Diagnosis. Maximum SVL 46.7 mm; eight or nine supralabials; 7–9 infralabials; ventral scales keeled; 5–7 continuous, pore-bearing precloacal scales with round pores; 22–27 paravertebral tubercles; body tubercles semi- linearly arranged, weakly present on flanks; tubercles present in lateral caudal furrows; no ventrolateral row of caudal tubercles; lateral row of caudal tubercles present; caudal tubercles not encircling tail; all subcaudals keeled, no enlarged median scale row; three postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no enlarged submetatarsal scales on first toe; 22–26 subdigital fourth toe lamellae; light- colored, vertebral stripe variably present (Tables 6,7).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 81 117 FIGURE 44. Cnemaspis temiah sp. nov. from Tanah Rata, Cameron Highlands, Pahang, Peninsular Malaysia. Upper left: adult female (LSUDPC 5677) in the dark color pattern phase. Upper right: hatchling (LSUDPC 5382). Middle right: adult male (LSUDPC 5668) in the light color pattern phase. Middle left: microhabitat at trail 10, Tana Rata, Cameron Highlands. Lower: type series of C. temiah sp. nov., adult male holotype (LSUHC 9110) on far left. Photographs by LLG.

Description of holotype. Adult female; SVL 38.6 mm; head oblong in dorsal profile, moderate in size (HL/ SVL 0.27), somewhat narrow (HW/SVL 0.18), flattened (HD/HL 0.39), distinct from neck; snout short (ES/HL 0.46), flat in lateral profile; postnasal region constricted medially, flat; scales of rostrum keeled, slightly raised, larger than similarly shaped scales on occiput; low, supraorbital ridges; weak frontorostral sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.21); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral slightly concave, dorsal 75% divided by longitudinal groove; rostral bordered posteriorly by supranasals and two smaller scales and laterally by first supralabials; 9R,8L raised supralabials of similar size; 9R,L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly;

82 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 118 bordered posteriorly by small, granular, postnasal scales; mental large, triangular, bordered posteriorly by four postmentals, outer two largest; gular scales raised, keeled; throat scales larger, raised, keeled. Body slender, not particularly long (AG/SVL 0.43); small, keeled, dorsal scales equal in size throughout body, intermixed with much larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles on flanks prominent; 26 paravertebral tubercles; pectoral scales raised, keeled, not elongate; abdominal scales slightly larger than dorsals, flat, keeled; no precloacal pores; forelimbs moderately long (FL/SVL 0.15), slender; dorsal scales of brachium raised, keeled; dorsal scales of forearm keeled, raised; ventral scales of brachium keeled, raised, juxtaposed; ventral scales of forearm smooth, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.20); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; toes increase in length from first to fourth with fourth being slightly longer than fifth; 25 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales raised anteriorly, weakly keeled, juxtaposed; middorsal and lateral caudal furrows present; no row of enlarged, median subcaudal scales; subcaudal scales keeled; caudal tubercles do not encircle tail; caudal tubercles present in lateral caudal furrow; no enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail. Color pattern in life (Fig. 44). Dorsal ground color light brown to yellowish; head, body (including flanks), and limbs overlain with irregularly shaped, blotched, dark markings, those in the paravertebral region somewhat paired and alternating with somewhat larger, yellowish marking; tail generally immaculate; ground color of all ventral surfaces beige, weak dark stippling on throat, pectoral region, limbs and tail. There is no sexual dimorphism in color pattern and the pattern lights considerably at night. Variation. Paratypes LSUHC 9160 and 9816 resemble the holotype in all aspects of coloration and pattern (Fig. 44). The other paratypes show differing degrees of vertebral stipping. LSUHC 9159 and 9739 have a wide, yellow vertebral stipe extending from occiput to onto the tail. LSUHC 9817 also bears a vertebral stripe but it is interrupted just posterior to the shoulder region. LSUHC 9818 has a vertebral stripe that extends only to a point midway down the body between the limb insertions. LSUHC 9159–60,9739, 9816 have broken tails. LSUHC 9816 is a male with a series of seven, contiguous, pore-bearing precloacal scales with round pores. Morphometric variation and variation in scalation are presented in Table 9.

TABLE 9. Meristic and mensural character states of the type series of Cnemaspis temiah sp. nov. w = weak; f = female; m = male; ant = anterior; and b = broke. Meristic abbreviations are listed in the Materials and Methods.

LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC 9110 9159 9160 9739 9816 9817 9818

holotype paratype paratype paratype paratype paratype paratype Supralabials 9 8 9 8 9 9 8 Infralabials 9 7 9 8 7 8 8 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 1 No. of precloacal pores / / / / 7 / / Precloacal pores continuous (1) or separated (0) / / / / 1 / / Precloacal pores elongate (1) or round (0) / / / / 0 / / No. of paravertebral tubercles 26 23 27 22 26 22 22 Tubercles linearly arranged (1) or more random (0) w 0 1 1 w 0 0 ...... continued on the next page

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 83 119 TABLE 9. (Continued) LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC 9110 9159 9160 9739 9816 9817 9818

holotype paratype paratype paratype paratype paratype paratype Tubercles present (1) or absent (0) on flanks 1 w w w w 0 w Caudal tubercles in lateral furrow (1) or not (0) 1 1 1 1 1 1 1 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 0 0 0 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 1 1 1 Caudal tubercles restricted to a single paravertebral 0 0 0 0 0 0 0 row on each side (1) or not (0) Subcaudals keeled (1) or not (0) 1 1 1 1 1 1 1 Single median row of keeled subcaudals (1) or not (0) 0 0 0 0 0 0 0 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 0 0 0 0 0 0 Postcloacal spurs / / / / 3 / / Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 0 0 No. of 4th toe lamellae 25 22 22 26 24 26 22 sex f f f f m f f SVL 38.6 38.4 34.1 34 31.1 35.3 33.3 TL 48.7 19.3 16.5b 10.2b 19.4 42.3 42.4 TW 2.5 2.7 2.5 2.7 2.5 2.4 2.6 FL 5.9 6.5 5.3 5.9 5.2 5.7 5.3 TBL 7.6 7.9 6.2 7.1 6.1 7.3 6.8 AG 16.8 16.2 15.7 14.8 13.8 14.3 14.3 HL 10.5 10.6 4 9.7 8.2 9.3 9.2 HW 7 6.8 3 5.9 5.7 6.2 6 HD 4.1 4.4 5.8 4.1 3.5 4 3.8 ED 2.3 2.2 1.9 2.1 2 2.1 2 EE 2.9 3.2 2.8 2.8 2.3 2.7 2.9 ES 4.9 5.1 4.3 4.8 3.9 4.4 5 EN 3.7 4 3.3 3.5 3.2 3.1 3.5 IO 2.4 2.4 2 2.2 2 2.4 2.6 EL 0.9 0.9 0.5 0.8 0.5 0.9 0.4 IN 1.2 0.8 0.8 0.9 0.8 0.9 1.1

Distribution. Cnemaspis temiah sp. nov. is known only from the Cameron Highlands plateau, Pahang, Peninsular Malaysia (Fig. 3). Natural History. Grismer (2011a) reported Cnemaspis temiah sp. nov. to be an inhabitant of hill dipterocarp forests occurring in the vicinity of 1,150 meters in elevation (Fig. 44). Unlike most Sundaic species of Cnemaspis that have a high affinity for rocky microhabitats, C. temiah sp. nov. occurs exclusively on vegetation. It is assumed this species is secretive and diurnal although it has never been observed during the day. Grismer (2011a) noted that all the lizards observed at night were sleeping on the trunks of trees or on the surfaces of leaves. Cnemaspis temiah

84 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 120 sp. nov. appears to be very localized in distribution as well. At Cameron Highlands, lizards are found regularly but only along one 200 m stretch of a certain trail. They are very rare elsewhere. Gravid females carrying two eggs have been found during March, April and October (Grismer 2011a), suggesting C. temiah sp. nov. may breed year- round. Etymology. The specific epithet temiah is an invariable noun in apposition in reference to the Temiah Tribe of Orang Asli people that are also endemic to the Cameron Highland region. Comparisons. Cnemaspis temiah sp. nov. is a member of the affinis group and was previously considered conspecific with C. flavolineata (see Grismer 2011a and references therein). The molecular analysis indicates that not only is C. temiah sp. nov. separate from C. flavolineata but may not even its closest relative (Fig. 2). Cnemaspis temiah sp. is separated from C. flavolineata by not having caudal tubercles that encircle the tail as opposed to having caudal tubercles encircling the tail anteriorly. It is further separated by lacking as opposed to having large, white, dorsal tubercles on the body and tail and having an uncorrected p-distance of 18.0% (Table 4). Cnemaspis temiah sp. nov. differs from species of the affinis group as follows. From all species of the affinis group except C. bayuensis, C. flavolineata, C. hangus sp. nov., C. selamatkanmerapoh, and C. stongensis sp. nov. C. temiah sp. nov. differs by lacking an ocellus in the shoulder region. From all species of the affinis group except C. harimau it differs by having as opposed to lacking caudal tubercles that encircle the tail at least anteriorly. Cnemaspis temiah sp. nov. differs from C. affinis, C. bayuensis, C. hangus sp. nov., C. selamatkanmerapoh, and C. stongensis sp. nov. and C. mcguirei by having fewer subdigital lamellae on the fourth toe (22–26 versus 27–35 collectively). From C. affinis, C. bayuensis, C. grismeri, C. harimau, and C. narathiwatensis it differs by having as opposed to lacking precloacal pores. Cnemaspis temiah sp. nov. differs further from C. harimau and C. selamatkanmerapoh in having 22–27 as opposed to 18–20 and 30 paravertebral tubercles, respectively. From C. affinis and C. bayuensis it differs in having as opposed to lacking tubercles in the lateral caudal furrows. It differs from C. shahruli by not having a yellow gular region and from C. grismeri and C. mcguirei by not having nearly immaculate white caudal bands and a wide, yellow, postscapular band. Relationships. The sister species relationship between Cnemapspis temiah sp. nov. and C. flavolineata is not supported and the two are separated by an uncorrected p-distance of 18.0% (Fig. 2; Table 4).

Cnemaspis narathiwatensis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Narathiwat Rock Gecko Fig. 45

Holotype. THNHM 1436. Type locality “Waeng District, Narathiwat Province, Thailand. Exact locality, collector, and date of collection unknown”. Diagnosis. Maximum SVL 53.2 mm; nine or 10 supralabials; 7–11 infralabials; keeled ventral scales; 3–6 discontinuous, pore-bearing precloacal scales with round pores; 28–34 paravertebral tubercles; body tubercles randomly arranged, present or absent on flanks; tubercles in lateral caudal furrows; ventrolateral row of caudal tubercles present; lateral row of caudal tubercles present; caudal tubercles do not encircle tail; subcaudals keeled; no enlarged, median row of subcaudals; 1–3 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe not enlarged; subtibials keeled; 24–30 subdigital fourth toe lamellae; white ocellus in shoulder region of males; light-colored vertebral stripe variably present; whitish to yellow bars on flanks; black and white caudal bands posteriorly (Tables 6,7). Color pattern (Fig. 45). Dorsal ground color of head, body, limbs and tail gray to brown; top of head mottled with yellow, bearing two dark, diffuse, postorbital stripes; lower postorbital stripe extending onto upper portion of forelimb; upper postorbital stripe wider, incomplete, extending onto occiput and nearly meeting opposing stripe; shoulder region dark and in males encloses a whitish ocellus composed of large tubercles; a series of light-colored bars on flanks which tend to fade posteriorly; small, dark, paired, paravertebral markings on trunk between forelimb insertion and base of tail alternating with large, round, light-colored paravertebral markings; ventrolateral tubercles on base of tail light-colored; limbs generally uniform light-brown to yellowish bearing small, randomly arranged, diffuse markings; ventral surfaces uniform beige with fine, dark stippling in each scale; throat darker; infralabials and mental light-yellow; subcaudal region bearing whitish rings. Distribution. Cnemaspis narathiwatensis is known only from the Thai-Malaysian border region from Hala-

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 85 121 Bala of the district of Waeng in Narathiwat Province and from Bang Lang National Park and Bannang Sata District, Yala Province, Thailand and the Belum-Temengor region of Perak in northern Peninsular Malaysia (Fig. 3). Natural history. Grismer et al (2010a) noted that lizards from Thailand were observed at night sheltering in rocky crevices between 200–500 m in elevation and suggested this may indicate this species is a diurnal, rocky, microhabitat specialist (Fig. 45). This was confirmed with findings of C. narathiwatensis in Belum-Temengor region of Peninsular Malaysia where lizards were found on granite boulders near a stream during the day but were not seen at night. Relationships. Cnemaspis narathiwatensis may be the sister species of C. shahruli (Fig. 2) also this relation is not statistically supported. Material examined. Thailand: Narathiwat Province, Waeng District THNHM 1338, 1436; Yala Province, Bannang Sata District, Bang Lang National Park THNHM 12435. These specimens represent the type series. Material examined since Grismer et al. (2010a): Malaysia: Perak, Belum-Temengor region LSUHC 11271–74.

FIGURE 45. Cnemaspis narathiwatensis from Sungai Enam, Belum, Perak, Peninsular Malaysia. Upper left: ventral view of an adult male and female (LSUDPC 7940). Upper right: adult female (LSUDPC 7009) in the dark color pattern phase. Lower right: adult male (LSUDPC 7011) in the dark color pattern phase. Lower left: granite boulder microhabitat at Sungai Enam. Photographs by ESHQ.

Cnemaspis hangus sp. nov. Bukit Hangus Rock Gecko Fig. 46

Holotype. Adult male (HC 00227) collected on 24 June 2008 by Chan Kin Onn at 1030 hrs from Bukit Hangus, Pahang, Peninsular Malaysia (04°16.142’N, 102°13.370’E) at 10 m in elevation. Paratypes. Adult female (HC 00225) has the same collection data as the holotype. Diagnosis. Cnemaspis hangus sp. nov. differs from all other Southeast Asia species of Cnemaspis in having the unique combination of adult males reaching 50.5 mm SVL, adult females reaching 47.0 mm SVL; nine supralabials; eight infralabials; ventrals keeled; no precloacal pores; moderately prominent dorsal tubercles; 22–24

86 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 122 paravertebral tubercles; dorsal body tubercles semi-randomly arranged; tuberculation weak on flanks; caudal tubercles not encircling tail; tubercles may be within lateral caudal furrows anteriorly only; lateral row of caudal tubercles present; ventrolateral caudal tubercles absent; subcaudals keeled; no enlarged, median subcaudal scale row; two postcloacal tubercles; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; and 27–34 subdigital lamellae on fourth toe (Tables 6,7). Cnemaspis hangus sp. nov. lacks the diagnostic color pattern characteristics of other species in the Peninsular clade. Description of holotype. Adult male; SVL 50.5 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.25), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.42), distinct from neck; snout short (ES/HL 0.48), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum weakly keeled, slightly raised, same size as similarly shaped scales on occiput; low, supraorbital ridges; moderate frontorostral sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.22); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral slightly concave, dorsal 75% divided by longitudinal groove; rostral bordered posteriorly by supranasals and one small, azygous scale and laterally by first supralabials; 9R,L raised supralabials of similar size; 8R,L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, bordered posteriorly by six small postmentals of similar size; gular scales raised, weakly keeled; throat scales larger, raised, keeled.

FIGURE 46. Cnemaspis hangus sp. nov. from Bukit Hangus, Pahang, Peninsular Malaysia. Upper left: adult male holotype (HC 0227) in the light color pattern phase. Lower left: ventral view of adult female (HC 0225) showing same coloration as adult males. Right: karst microhabitat at Bukit Hangus. Photographs by CKO.

Body slender, elongate; small, keeled, dorsal scales equal in size throughout body, intermixed with larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles on flanks sparse, moderate in size; 22 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales slightly larger than dorsals; no precloacal pores; forelimbs moderately long, slender; dorsal scales of brachium raised, keeled; dorsal scales of forearm keeled, raised; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm weakly raised, juxtaposed; palmar scales

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 87 123 smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing present; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh weakly keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally but wider distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing present; toes increase in length from first to fourth with fourth being slightly longer than fifth; 34 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; caudal scales flat anteriorly, weakly keeled, juxtaposed; deep middorsal and lateral furrows; no enlarged, median subcaudal scales; subcaudal scales keeled; no median row of enlarged keeled subcaudal scales; caudal tubercles do not encircle tail; caudal tubercles may or may not be present in lateral furrow anteriorly; one enlarged postcloacal tubercle on lateral surface of hemipenal swellings at base of tail. Color pattern in life (Fig. 46). Dorsal ground color dark, ashy grey; head and body overlain with irregularly shaped, indistinct darker blotchs; light yellowish, medial blotch on occiput following three smaller dark spots; rostrum bearing yellowish spots; single, faded, dark, postorbital stripe extending to base of occiput; paravertebral, faded yellowish markings on extending to base of tail alternating with faded irregularly shapred, smaller, dark markings; transversely elongate, yellowish markings on flanks alternating with darker spots; tail regenerated, unicolor gray; irregularly shaped, small, dark and light markings on limbs; dark and light diffuse bands encircling digits; ventral surfaces dark grey. There is no sexual dimorphism in color pattern and the pattern lightens considerably at night. Variation. The paratype (HC 0225) approaches the holotype in coloration and pattern except that its dorsal pattern is bolder, its ventral surfaces are slightly lighter, and most of its tail is original and bears a faded banding pattern. Morphometric variation and variation in scalation are presented in Table 10.

TABLE 10. Meristic and mensural character states of the type series of Cnemaspis hangus sp. nov. w = weak; f = female; and m = male. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right.

HC HC 227 225 holotype paratype SVL (mm) 50.5 47 Supralabials 9 9 Infralabials 8 8 Ventral scales keeled (1) or not (0) 1 1 No. of precloacal pores 0 / Precloacal pores continuous (1) or separated (0) / / Precloacal pores elongate (1) or round (0) / / No. of paravertebral tubercles 22 24 Tubercles linearly arranged (1) or more random (0) 0 0 Tubercles present (1) or absent (0) on flanks w w Caudal tubercles in lateral furrow (1) or not (0) ant 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 0 0 Lateral caudal tubercle row present (1) or absent (0) 1 1 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 0 Subcaudals keeled (1) or not (0) 1 1 ...... continued on the next page

88 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 124 TABLE 10. (Continued) HC HC 227 225 holotype paratype Single median row of keeled subcaudals (1) or not (0) 0 0 Caudal tubercles encircle tail (1) or not (0) 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 0 Postcloacal spurs 2 / Enlarged femoral scales present (1) or absent (0) 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 Subtibial scales keeled (1) or not (0) 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 No. of 4th toe lamellae 34 27 Sex m f TL 16.1/44.4 28.3/21.3 TW 5.3 4.5 FL 8.8 8.4 TBL 10.3 9.8 AG 21.7 20.4 HL 12.7 13.1 HW 8.7 8.6 HD 5.3 5.5 ED 2.8 3.2 EE 3.7 3.9 ES 6.1 5.5 EN 4.6 4.2 IO 2.7 2.3 EL 1.5 1.2 IN 1 /

Distribution. Cnemaspis hangus sp. nov. is known only from Bukit Hangus, Pahang, Peninsular Malaysia (Fig. 3). Natural History. Lizards were seen abroad during the day on karst boulders and walls within a lowland dipterocarp forest and on boulders near the entrances of cave openings (Fig. 46). All were wary and quick to take cover in crevices and on the backsides of boulders at the slightest provocation. Multiple lizards were observed to occupy the same boulder and no lizards were seen at night indicating that C. hangus sp. nov. is diurnal. HC 0225 was a gravid female carrying two eggs indicating that reproduction takes place in June. Etymology. The specific epithet hangus is an invariable noun in apposition in reference to the Malay word “hangus” which means to burn or scorch and refers to this species’ overall burnt appearance in the dark color pattern phase. Comparisons. Cnemaspis hangus sp. nov. is a member of the affinis group within which it is the sister species to C. selamatkanmerapoh. Cnemaspis hangus sp. nov. differs from C. selamatkanmerapoh in having a larger maximum larger SVL (50.5 mm versus 43.4 mm); lacking precloacal pores as opposed to having pores; having fewer paravertebral tubercles (22–24 versus 30); and semi-randomly arranged versus linearly arranged, dorsal tubercles. Cnemaspis hangus sp. nov. and C. selamatkanmerapoh bear a 3.6% sequence divergence from one another as well (Table 4). It differs from C. pseudomcguirei, C. harimau, and C. shahruli by having a greater maximum SVL (50.5 mm versus 36.5–43.2) and being considerably smaller than C. mcguirei (maximum SVL 65.0 mm). It differs from all species of the affinis group except C. shahruli in lacking as opposed to having precloacal

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 89 125 pores. From C. mcguirei, C. grismeri and C. narathiwatensis in having fewer paravertebral tubercles (22–24 versus 23–34) and from C. harimau and C. shahruli in have more precloacal pores (11–23). It can be further separated from all species of the affinis group except C. affinis, C. bayuensis and C. selamatkanmerapoh by lacking tubercles in the lateral caudal furrows. From C. grismeri and C. narathiwatensis it differs in lacking as opposed to having a row of ventrolateral caudal tubercles. Having 27–34 subdigital lamellae on the fourth toe separates if from C. pseudomcguirei, C. flavolineata and C. temiah sp. nov. (23–26). Relationships. Cnemaspis hangus sp. nov. is the sister species of C. selamatkanmerapoh (Fig. 2).

Cnemaspis selamatkanmerapoh Grismer, Wood, Mohamed, Chan, Heinz, Sumarli, Chan & Loredo, 2013a Merapoh Rock Gecko Fig. 47

Holotype. Adult male (LSUHC 11016) “collected on 23 June 2013 by L. Lee Grismer at 2200 hrs at 23 m from Gua Gunting, Merapoh, Pahang, Peninsular Malaysia (4°42.069 N, 101°58.512 E)”. Diagnosis. Maximum SVL 43.4 mm SVL; 10 supralabials; nine or 10 infralabials; keeled ventrals; at least one, round precloacal pore in males; 30 paravertebral tubercles; dorsal body tubercles semi-randomly arranged; tuberculation weak on flanks; caudal tubercles not encircling tail; lateral caudal tubercles not within lateral caudal furrows; ventrolateral caudal tubercles absent; subcaudals keeled; no enlarged, median subcaudal scale row; three postcloacal tubercles; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; and 31–33 subdigital lamellae on fourth toe (Tables 6,7). Cnemaspis selamatkanmerapoh sp. nov. lack diagnostic color pattern characteristics. Color pattern in life (Fig. 47). Dorsal ground color grey; top of head bearing small dark spots; thin, dark postorbital stripes meeting medially on occiput and turning anteriorly; rostrum and supralabial region greenish; paired, light colored, paravertebral, blotches extend from nape to base of tail where they transform into light colored caudal bands, blotches united on nape and shoulder region into a single blotch; flanks bearing dark mottling and yellowish spots; limbs darkly mottled with a faint banding pattern; overall color of venter unicolor beige with all scales bearing black stippling. There is no sexual dimorphism in color pattern and coloration lightens considerably at night. Distribution. Cnemaspis selamatkanmerapoh is known only from the type locality of Gua Gunting, Merapoh, Pahang, Peninsular Malaysia (Fig. 2). Grismer et al. (2013a) reported finding eggs on the connected karst outcrop Gua Goyang. Natural History. Grismer et al. (2013a) noted that Cnemaspis selamatkanmerapoh is a lowland, diurnal species found only on karst substrate. Specimens were observed approximately 1 m above the ground along the perimeter of an extensive karst system surrounded by a limestone forest (Fig. 47) and lizards often position themselves adjacent to cracks into which they can escape if threatened. A gravid female carrying two eggs was collected during June. Relationships. Cnemaspis selamatkanmerapoh is the sister species of C. hangus sp. nov. (Fig. 2). Material examined. Malaysia: Pahang; Mearpoh, Gua Gunting LSUHC 11015–16 (type series).

Cnemaspis bayuensis Grismer, Grismer, Wood & Chan, 2008 Kampung Bayu Rock Gecko Fig. 48

Holotype. ZRC 2.6759. Type locality: “Gua Bayu, Kelantan, Peninsular Malaysia (05°05.650 N, 102°13.265 E)” at 120 m in elevation. Diagnosis. Maximum SVL 46.1 mm; nine or 10 supralabials; eight or nine infralabials; ventral scales keeled; 5–9 discontinuous, pore-bearing precloacal scales with round pores; 23–30 paravertebral tubercles; tubercles not linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral caudal row present; caudal tubercles not restricted to a single paravertebral row nor encircling tail; all subcaudals keeled, no enlarged median scacle row; two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; subtibials keeled; no submetatarsal scales on first toe; 27–32 subdigital fourth toe lamellae; whitish bars on flanks (Tables 6,7).

90 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 126 FIGURE 47. Upper: adult male holotype (LSUHC 11016) Cnemaspis selamatkanmerapoh in the dark color pattern phase. Lower: microhabitat at Gua Goyang, Merapoh, Pahang, Peninsular Malaysia. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 91 127 FIGURE 48. Cnemaspis bayuensis from Gua Bayu, Kelantan, Peninsular Malaysia. Left: karst microhabitat of C. bayuensis near Kampung Bayu. Upper right: adult male (LSUDPC 4417) in the dark color pattern phase. Lower right: adult female (LSUDPC 4420) in the light color pattern phase. Photographs by LLG.

Color pattern in life (Fig. 48). Ground color of dorsal surfaces of head, body, limbs, and tail brown; head and body overlain with irregularly shaped, dark spots; light markings on occiput; a single, thin, dark, postorbital stripe extends onto nape does not contact medially with an opposing postorbital stripe; dark, anteriorly projecting, triangular marking between opposing postorbital stripes; white, paravertebral markings on nape followed by distinct, white, alternating, paravertebral blotches extending to base of tail; transversely elongate, distinct, white bars on flanks; dark blotches often on body; diffuse brown and mottled white bands encircle tail; dark caudal banding sometimes present; irregularly shaped, dark and light markings on limbs; dark and light, diffuse bands encircle digits; gular region bears a faint, brown, reticulate pattern; ventral surfaces of body and limbs dull beige, immaculate and darkened laterally; subcaudal region suffused with dark pigment. Distribution. Cnemaspis bayuensis is known only from the type locality of Gua Bayu, Kelantan, Peninsular Malaysia (Grismer et al. 2008b: Fig. 3) but is likely to be found throughout the nearby karst region. Natural history. Grismer et al. (2008b) noted that Cnemaspis bayuensis is a saxicolous species restricted to the lowland karst outcroppings (Fig. 48) of the Gua Bayu region surrounding the village of Bayu. Lizards are diurnal and occur along the periphery of the karst formations in cracks as well as on cave walls and ceilings as high as 5 m above the ground. Lizards have not been found deep within the cave systems. This species is adept at matching the color of its substrate be it the light colored cave walls or the dark, lichen-colored isolated karst boulders scattered along the periphery of the karst towers. Females carrying two eggs have been observed during mid-June. Relationships. Cnemaspis bayuensis is the sister species of C. stongensis sp. nov. (Fig. 2). Material examined. Malaysia: Kelantan, Gua Bayu ZRC 2.6759–61, LSUHC 9073 (type series).

92 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 128 Cnemaspis stongensis sp. nov. Gunung Stong Rock Gecko Fig. 49

Holotype. Adult male LSUHC 11089 collected on 26 June 2013 by Chan Kin Onn, L. Lee Grismer and Jacob A. Chan at 1030 hrs at 10 m from Kem Baha, Gunung Stong, Kelantan, Peninsular Malaysia (5°20.465 N, 101°58.001 E) at 461 m elevation. Paratypes. Adult males LSUHC 11091, 11093–94, 11100, 11139 and adult female LSUHC 11092 has the same collection data as the holotype. Diagnosis. Cnemaspis stongensis sp. nov. differs from all other Southeast Asia species of Cnemaspis in having the unique combination of adult males reaching 49.3 mm SVL, adult females reaching 48.4 mm SVL; 8–11 supralabials; 8–10 infralabials; ventrals keeled; 5–8, continguous, pore-bearing precloacal scales with round pores; moderately prominent dorsal tubercles; 26–33 paravertebral tubercles; dorsal body tubercles generally randomly arranged; tubercles present on flanks; caudal tubercles not encircling tail; lateral caudal tubercles usually within lateral caudal furrows anteriorly only; ventrolateral caudal tubercles may or may not be present anteriorly only; lateral row of caudal tubercles present anteriorly only; subcaudals keeled; no enlarged, median subcaudal scale row; two or three postcloacal tubercles; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; and 28–32 subdigital lamellae on fourth toe. These differences are summarized across all Southeast Asian species in Table 6. Cnemaspis stongensis sp. nov. lacks diagnostic color pattern characteristics. Description of holotype. Adult male; SVL 49.3 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.25), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.43), distinct from neck; snout short (ES/HL 0.49), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum keeled, raised, slightly larger than similarly shaped scales on occiput; low, supraorbital ridges; moderate frontorostral sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.23); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave, dorsal 90% divided by longitudinal groove; rostral bordered posteriorly by two small supranasals and two large scales between the supranasals and laterally by first supralabials; 9R,L raised supralabials of similar size; 8R, 9L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, bordered posteriorly by four small postmentals of similar size; gular scales raised, keeled; throat scales same size, raised, keeled. Body slender, elongate (AG/SVL 0.43); small, keeled, dorsal scales equal in size throughout body, with intermixed larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles on flanks sparse, moderate in size; 32 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales slightly larger than dorsals; eight pore-bearing precloacal scales with round pores in a chevron pattern separated on the left side of the chevron by a single non- pore-bearing scale; forelimbs moderately long, slender (FL/SVL 0.18); dorsal scales of brachium raised, keeled; dorsal scales of forearm keeled, raised; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm raised, juxtaposed; palmar scales raised, smooth, juxtaposed; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wider; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.21); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales raised, smooth, juxtaposed; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally but wider distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wider; interdigital webbing absent; toes increase in length from first to fourth with fourth being slightly longer than fifth; 32 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; caudal scales flat anteriorly, weakly keeled, juxtaposed; middorsal and lateral furrows; no enlarged, median subcaudal scales; subcaudal scales keeled; caudal tubercles present in lateral furrow anteriorly; two enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 93 129 FIGURE 49. Cnemaspis stongensis sp. nov. from Kem Baha, Gunung Stong, Peninsular Malaysia. Upper left: adult male holotype (LSUHC 11091) in dark color pattern phase. Upper right: ventral view of the holotype and a gravid female paratype (LSUHC 11091 upper and 11100 lower, respectively). Middle left: adult female paratype (LSUHC 11100) in dark color pattern phase. Middle right: granite boulder microhabitat at Kem Baha, Gunung Stong, Peninsular Malaysia. Lower: type series of C. stongensis sp. nov. Photographs by LLG.

Color pattern in life (Fig. 49). Dorsal ground color of head, limbs, and body grey; black, irregular striping on snout; black, transverse, azygous marking on top of head; black, postorbital stripe extending onto nape; black, medial, teardrop-shaped marking on nape; a series of seven, black, paravertebral markings extending from shoulder region to base of tail; similar black markings on flanks; dull-white, butterfly-shaped, paravertebral markings alternating with black, paravertebral markings and becoming completely separated posteriorly; dull-white,

94 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 130 transverse bars on flanks; light-colored, caudal bands, anterior two yellowish and posterior seven white; light caudal bands alternate with black caudal bands; limbs bearing irregularly shaped, whitish bands; ventral surfaces of head, throat, pectoral, and abdominal regions, and limbs grey, gular region and abdomen slightly darker; caudal bands encircle tail but are less vivid in subcaudal region. Variation. The paratypes closely resemble the holotype in coloration and pattern (Fig. 49). The dorsal pattern of LSUHC 11094 and 11139 appears slightly more speckled overall and LSUHC 11100 has a faint, wide, light- colored, vertebral stripe. Morphometric variation and variation in scalation are presented in Table 11. Distribution. Cnemaspis stongensis sp. nov. is known only from the type locality and surrounding areas on Gunung Stong, Kelantan, Peninsular Malaysia (Fig. 3). Natural History. Lizards were observed to be active only at night on large granite boulders (Fig. 49) ranging in elevation from 50 m at the base of Gunung Stong at Hutan Lipur Jelawang in lowland dipterocarp forest up to the type locality of Kem Baha at 461 m in hill dipterocarp forest. The majority of specimens were found in the vicinity of water but were not necessarily restricted to these areas. Within their microhabitat, lizards were most commonly seen on the undersides or on the lower sections of boulders (Fig. 49) making escape much easier. We expect this species extends to higher elevations on Gunung Stong above Kem Baha.

TABLE 11. Meristic and mensural character states of the type series of Cnemaspis stongensis sp. nov. w = weak; f = female; m = male; ant = anterior; and r = regenerated. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right.

LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC

11089 11091 11092 11093 11094 11100 11139

holotype paratype paratype paratype paratype paratype paratype

Supralabials 9 8 10 8 9 10 11 Infralabials 8 8 9 8 10 7 9 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 1 No. of precloacal pores 8 6 / 7 5 5 5 Precloacal pores continuous (1) or separated (0) 1 1 / 1 1 1 1 Precloacal pores elongate (1) or round (0) 0 0 / 0 0 0 0 No. of paravertebral tubercles 32 29 30 33 29 28 26 Tubercles linearly arranged (1) or more random (0) 0 0 0 0 0 0 0 Tubercles present (1) or absent (0) on flanks 1 1 1 1 1 1 1 Caudal tubercles in lateral furrow (1) or not (0) ant ant ant 0 ant / ant Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 0 1 1 / 1 Lateral caudal tubercle row present (1) or absent (0) ant ant ant ant ant / ant Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 0 0 0 0 / 0 Subcaudals keeled (1) or not (0) 1 1 1 1 1 / 1 Single median row of keeled subcaudals (1) or not (0) 0 0 0 0 0 / 0 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 0 / 0 Enlarged median subcaudal scale row (1) or not (0) 0 0 0 0 0 / 0 Postcloacal spurs 2 2 / 2 2 3 2 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 1 1 ...... continued on the next page

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 95 131 TABLE 11. (Continued)

LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC

11089 11091 11092 11093 11094 11100 11139

holotype paratype paratype paratype paratype paratype paratype

Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 0 0 No. of 4th toe lamellae 32 28 28 29 31 31 31 Sex m m f m m m m SVL 49.3 48.4 46.2 44.3 43.9 48.5 47.2 TL 68.9 67.6 62.1 49.5/11.1 61.5 42.3r 38.1/,19.1 TW 4.8 5.3 5.1 4.4 5.4 4.7 4.6 FL 8.8 8.3 7.9 8.4 8.2 8.3 9.1 TBL 10.3 10.1 9.4 9.9 9.9 10.9 10.4 AG 21.2 20.1 21.1 18.4 19.7 20.9 21.1 HL 12.1 13.1 12.2 12.5 11.9 12.9 13.1 HW 8.4 8.3 7.9 7.5 7.7 8.3 7.9 HD 5.2 5.5 4.5 4.8 4.7 5.3 5.3 ED 2.8 2.7 2.3 2.4 2.3 2.6 2.5 EE 4.1 4.1 4.1 3.5 3.3 3.9 3.7 ES 5.9 6.3 5.9 5.5 5.6 5.9 6.1 EN 4.6 4.7 4.4 4.4 4.2 4.5 4.6 IO 2.7 2.9 2.6 2.7 2.7 2.9 2.8 EL 0.9 1.2 0.9 1.1 0.9 0.7 0.9 IN 1.3 1.1 1.1 1.2 1.2 1.3 1.2

Etymology. The specific epithet stongensis is an adjective in reference to Gunung (mountain) Stong on which the type locality of Kem Baha is located. Comparisons. Cnemaspis stongensis sp. nov. is a member of the affinis group within which it is the sister species of C. bayuensis. Cnemaspis stongensis sp. nov. differs from C. bayuensis in having caudal tubercles anteriorly in the lateral caudal furrow as opposed to lacking them; having a lateral row of caudal tubercles present anteriorly as opposed to throughout the length of the tail; and lacking distinct black and white caudal bands as opposed to having them. However, these two species bear only a 2.2% sequence divergence from one another (Table 4). Cnemaspis stongensis sp. nov. differs from C. pseudomcguirei, C. harimau, and C. shahruli by having a greater maximum SVL (49.3 mm versus 36.5–43.2) and being considerably smaller than C. mcguirei (maximum SVL 65.0 mm). It differs from C. shahruli and C. hangus sp. nov. in having as opposed to lacking pre-cloacal pores. From C. harimau, C. shahruli, C. flavolineata, C. temiah sp. nov. and C. hangus sp. nov. it differs in having more paravertebral tubercles (26–33 versus 18–27). Cnemaspis stongensis sp. nov. can be further separated from all species of the affinis group except C. affinis, C. hangus sp. nov., C. selamatkanmerapoh, and C. bayuensis by generally lacking tubercles in the lateral caudal furrows. It is seaparated from C. harimau and C. temiah sp. nov. by not having caudal tubercles that encircle the tail. Having 28–32 subdigital lamellae on the fourth toe separates it from C. pseudomcguirei, C. flavolineata, and C. temiah sp. nov. Comments. The relatively close morphological and genetic similarity between Cnemaspis stongensis sp. nov. and C. bayuensis is surprising in that these sister species are separated by at least 85 km of uninhabital terrane and the former is a granite boulder specialist and the latter is a karst specialist. Relationships. Cnemaspis stongensis sp. nov. is the sister species of C. bayuensis (Fig. 2). Additional material examined. Malaysia: Kelantan; Gunung Stong, Kem Baha LSUHC 11090, 11095, 11138.

96 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 132 Southern Sunda clade

The Southern Sunda clade is a strongly supported lineage that contains the 12 southernmost species of Cnemaspis (Fig. 2) 10 from which we had molecular data. This clade ranges from southern Peninsular Malaysia and Singapore eastward through the Seribuat, Anambas, and Natuna archipelagos, to at least west-central and northern Borneo (Figs. 1,3). It forms a polytomy comprised of three well-supported lineages: the insular endemic C. limi Das & Grismer from Tioman and Tulai islands in the Seribuat Archipelago; the sister species C. nigridia (Smith) and C. paripari Grismer & Chan from northwestern Borneo referred to here as the nigridia group; and the kendallii group containing the remaining seven species, C. kendallii sensu stricto (Gray) from East Malaysia and Indonesia, C. sundainsula sp. nov. from Natuna Besar Island of the Natuna Archipelago, Indonesia and a lineage containing the Peninsular Malaysian C. pemanggilensis Grismer, Grismer & Das from Pulau Pemanggil, C. baueri Das & Grismer from Aur and Dayang islands, C. mumpuniae sp. nov. from Natuna Besar, Indonesia, C. bidongensis Grismer, Wood, Amirrudin, Sumarli, Vazquez, Ismail, Lukman, Nance, Muhammad, Mohaamad, Syed, Kuss, Murdoch & Cobos from Pulau Bidong and C. peninsularis sp. nov. from Peninsular Malaysia, Singapore, and the Seribuat Archipelago (Figs. 2,4). The relationship of these species is further supported in that they are the only Cnemaspis in the Southern Sunda clade that lack precloacal pores (Fig. 5) which is considered derived in that precloacal pores are widespread throughout the Gekkonidae (Kluge 1987). In the molecular analysis, there is an unresolved polytomy among these species (Fig. 2), however, the morphological analysis supports the monophyly of C. kendallii, C. pemanggillensis, C. baueri, C. mumpuniae sp. nov., C. bidongensis and C. peninsularis sp. nov. to the exclusion of C. sundainsula in that they are the only species of Cnemaspis in the Southern Sunda clade that have caudal tubercles encircling the tail (Fig. 5). Furthermore, the monphyly of C. baueri, C. mumpuniae sp. nov., C. bidongensis and C. peninsularis sp. nov., is supported in that they are the only species (with the exception of some individuals of C. tucdupensis from the chnathaburiensis group) wherein the posterior portion of the original tail is black (Fig. 5).

Cnemaspis limi Das & Grismer, 2003 Lim’s Rock Gecko Fig. 50

Cnemaspis sp. Hendrickson 1966:56 Cnemaspis nigridius Manthey & Grossmann 1997:214; Lim & Lim 1999:142 Cnemaspis cf. nigridia Chan-ard et al. 1999:104

Holotype. ZRC 2.5289. Type locality: “Gua Tengkuk Air, Gunung Kajang, adult male …collected from Pulau Tioman [Pahang] (02°50’ N, 104°09’ E), West Malaysia” at 980 m in elevation. Diagnosis. Maximum SVL 88.2 mm; 8–12 supralabials; 7–10 infralabials; ventral scales weakly keeled; no precloacal pores; 25–35 paravertebral tubercles; body tubercles randomly arranged, weakly present on flanks, absent from lateral caudal furrows; no ventrolateral caudal tubercles; lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals smooth bearing a median, weakly enlarged, scale row; one or two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials weakly keeled; 29–36 subdigital fourth toe lamellae; large, black, round spots on nape and anterior portion of body; dorsal caudal tubercles white; at least posterior one-half of subcaudal region white (Tables 6,7). Color pattern in life (Fig. 50). Dorsal ground color of head, body, limbs, and tail dark brown; thin yellow reticulum on top of head and body; a dark, upper, postorbital stripe extends onto nape; a dark, lower, postorbital stripe extends onto shoulder region; a medial, dark marking on nape; 5–7 dark, paravertebral spots occur on back and flanked by diffuse, dark blotches on sides; faint, dark mottling on hind limbs and more on forelimbs; diffuse, dark bands on tail; anterior two-thirds of tail encircled by large, white to cream-colored tubercles; ventral surfaces of head, lateral sections of belly, limbs, and anterior one-third of tail dull-brown, immaculate; ventral surfaces of belly beige. At night the brown ground color of the dorsum fades to white, accenting the black body spots and the yellow reticulum. Distribution. Cnemaspis limi is known only from Pulau Tioman, Pahang and the adjacent island of Tulai, Johor in the Seribuat Archipelago of Peninsular Malaysia (Grismer 2011a,b; Fig. 4).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 97 133 Natural History. According to Grismer (2011a), Cnemaspis limi is a saxicolous gecko inhabiting primary and secondary coastal vegetation in lowland and hill dipterocarp forests (Fig. 50) from sea level to the summit of Gunung Kajang at 1,026 m in elevation. Lizards are found almost exclusively on large granite boulders where, during the day, they are active on vertical, shaded surfaces, within crevices, and within the cave-like situations formed by the aggregations of boulders piled on top of one another. In the latter microhabitat, densities can be surprisingly high. Lizards usually sit facing head-down or upside down and run to the base of the rock to escape. At night, however, C. limi is less active and almost exclusively restricted to cave-like situations. This is especially true for lizards near the summit of Gunung Kajang. Bullock (1966) found the larvae of butterflies, grasshoppers, beetles, and pieces of ants in the stomachs of lizards he examined, indicating C. limi does not feed on the ground like C. peninsularis sp. nov. with which it is sympatric. Grismer (2011a) reported females carrying one or two eggs having been observed during July and September and eggs stuck to the undersides of rocks during April indicating C. limi breeds throughout the year. Relationships. Cnemaspis limi is a basal lineage in a tritomy composing the Southern Sunda clade (Fig. 2).

FIGURE 50. Cnemaspis limi from Pulau Tioman, Pahang, Peninsular Malaysia. Upper left: adult male (LSUDPC 6185) in the light color pattern phase. Upper right: adult female (LSUDPC 5623) in the dark color pattern phase. Lower: granite boulder microhabitat on the Tekek-Juara Trail, Pulau Tioman. Photographs by LLG.

Material examined. Peninsular Malaysia: Pahang, Pulau Tioman ZRC 2.5289–90, 2.3504–06 (type series). Material examined since Das and Grismer (2003): Peninsular Malaysia: Johor: Pulau Tulai LSUHC 5053; Pahang, Pulau Tioman LSUHC 3801–02, 3859, 3888, 3902, 3904, 4410, 4425, 4480–82, 4485–88, 4563–64, 4596, 5424, 5441, 5510, 5515, 5518, 5521, 6203, 6206–07, 6210, 6212, 6267, 8035.

98 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 134 Nigridia group. The nigridia group contains two very dissimilar species from extreme northwestern Borneo; the large, nocturnal, granite boulder-dwelling Cnemaspis nigridia and the much smaller, diurnal, karst-dwelling species C. paripari (Fig. 2). The monophyly of this group is further supported in that these are the only species in the Southern Sunda clade to have enlarged scales beneath the first metatarsals. This character state occurs in the two species of the Ca Mau clade, four of 22 species of the Northern Sunda clade and does not occur in the Pattani clade (Table 6). Therefore, we consider this character state to be derived. The nigridia group is diagnosed as having a maximum SVL of 50.7–75.5 mm; 10–13 supralabials; 9–11 infralabials; weakly to strongly keeled ventral scales; 2–16 contiguous, pore-bearing precloacal scales with round pores; 26–43 paravertebral tubercles; randomly to occasionally weakly aligned dorsal tubercles on the body; no tubercles on flanks or in the lateral caudal furrows; caudal tubercles do not encircle tail; lateral row of caudal tubercles present; subcaudals keeled and bearing a median row of enlarged, keeled scales; 2–4 postcloacal tubercles on either side of the base of the tail; no enlarged femoral or subtibial scales; subtibials keeled; submetatarsal scales of first toe enlarged; and 26–31 lamellae beneath the fourth toe.

FIGURE 51. Cnemaspis nigridia from Gunung Gading, Sarawak, East Malaysia. Upper left: adult female (LSUDPC 4882) in dark color pattern phase. Upper right: adult male (LSUDPC 4344) in light color pattern phase. Lower: granite boulder microhabitat at Gunung Gading. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 99 135 Cnemaspis nigridia (Smith, 1925) Black Rock Gecko Fig. 51

Heteronota Kendallii Gray, 1845:174 (in part) Gonatodes affinis Shelford, 1901:49 Gonatodes nigridius Smith, 1925:22 Cnemaspis nigridius Brongersma, 1934:165; Manthey & Grossmann, 1997:214; Cox, van Dijk, Nabhitabhata & Thirakhupt, 1998:90; Auliya 2006:180 Cnemaspis cf. nigridia Chan-ard, Grossmann, Gumprecht & Schulz, 1999:104

Holotype. BM 1946.8.22.90. Type locality: “Mt. Gadin” = Gunung Gading, Sarawak, East Malaysia (01°41.16 N, 109°50.35 E) at approximately 100 m in elevation. Diagnosis. Maximum SVL 75.5 mm; 10 or 11 supralabials; 9–11 infralabials; ventral scales weakly to moderately keeled; 10–16 contiguous, pore-bearing, precloacal scales with round pores; 39–43 paravertebral tubercles; tubercles semi-linear to randomly arranged, absent from flanks and lateral caudal furrows; ventrolateral caudal tubercles present; lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals keeled but bearing an enlarged median row of smooth scales; 2–4 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe enlarged; subtibials keeled; 26–29 subdigital fourth toe lamellae; a pair of large, round, black spots in shoulder region; yellow to white bars on flanks; wide, black and dull-yellow caudal bands (Tables 6,7). Color pattern in life (Fig. 51). Dorsal coloration of head, body and tail dark brown overlain with large, oval, black spots on anterior portion of body and nape; cream- colored, vertebral markings on nape and dorsum; thin, semi-transversely oriented, yellow bands on body and flanks; limbs bearing wide, dark and yellowish alternating faint bands; yellow caudal bands containing dark pigmentation; all ventral surfaces dark-gray. Distribution. Cnemaspis nigridia is known from the type locality of Gunung Gading as well as Gunung Pueh, Gunung Beremput, and the Bau Limestone Area (Smith 1925; Naming & Das 2004; Das 2006), Sarawak, East Malaysia (Fig. 4). Natural History. We have observed Cnemaspis nigridia only at night on the surfaces of large granite boulders in the vicinity of streams within old lowland secondary forest in forest (Fig. 51). Surveys during the day resulted in finding only C. kendallii in these areas. In all the locations from which this species has been reported except the Bau Limestone Area (which may be erroneous, see below) it is restricted to granite boulders. Remarks. Naming & Das (2004) report Cnemaspis nigridia as occurring on karst formations in the Bau Limestone Area. However, it is not clear if specimens were actually examined as there is no character support for their identification or voucher specimens listed. Only the habitat generalists C. kendallii and C. peninsularis sp. nov. and C. flavigaster occur on both karst and granite and we suspect the Bau population may not be C. nigridia but the karst-adapted C. paripari which occurs in that region (Grismer & Chan 2009). Relationships. Cnemaspis nigridia is the sister species of C. paripari from northwestern Borneo (Fig. 2). Material examined. East Malaysia: Sarawak, Gunung Gading LSUHC 9167–70.

Cnemaspis paripari Grismer & Chan, 2009 Fairy Rock Gecko Figs. 52, 53

Holotype. ZRC 2.6812. Type locality: “Gua Pari-pari, Bau District, Sarawak, [East] Malaysia (01°22.867 N, 110°07.164 E)” at approximately 30 m inelevation. Diagnosis. Maximum SVL 50.7 mm; 12 or 13 supralabials; 10 or 11 infralabials; ventral scales keeled; 2–6, discontinuous, pore-bearing, precloacal scales with round pores; 26–31 paravertebral tubercles; body tubercles randomly arranged, absent on flanks and from lateral caudal furrows; ventrolateral caudal tubercles absent; lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals keeled but bearing an enlarged median row of smooth scales; two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe enlarged; subtibials keeled; 26–31 subdigital fourth toe lamellae; head, limbs, and regenerated tail yellow in males; posterior one-half of original tail white in males (Tables 6,7).

100 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 136 FIGURE 52. Adult male Cnemaspis paripari (LSUDPC 4920) from Gua Pari-pari, Sarawak, East Malaysia in the dark color pattern phase. Photograph by LLG.

Color pattern in life (Figs. 52, 53). Males: dorsal ground color of body yellowish brown; limbs yellow; head (especially snout) bright yellow bearing small, irregularly shaped, brownish, occipital flecks and a faint, brownish, postorbital stripe; ground color of nape and shoulder region gray bearing paravertebral, irregularly shaped, black blotches; incomplete, transverse, yellow bands between forelimb and hind limb insertions; smaller, scattered, dark spots between bands; limbs generally immaculate; anterior one-half of tail gray bearing faint, dark bands; posterior one-half of tail immaculate, white; all ventral surfaces gray except for posterior half of tail which is white and beige; regenerated tail bright yellow. Females: adult females have an overall brown ground color, lack a yellow head and a yellow or white tail; yellow banding on body faint; dark blotching pattern on nape; tail brown at base, gradually turning to gray posteriorly; weakly banded. Distribution. Cnemaspis paripari is known only from the karst formations that extend approximately 4.2 km from Gua Angin to Gua Pari-pari within the Bau Limestone Area, Sarawak, Malaysia (Fig. 4). Natural History. According to Grismer and Chan (2009), Cnemaspis paripari is a diurnal, lowland, saxicolous species that appears to be restricted to the karst outcroppings extending from Gua Angin to Gua Pari- pari that are surrounded by lowland dipterocarp forest (Fig. 53). They reported seeing several specimens around and slightly within the openings of caves where light could still penetrate as well as on rocks along the periphery of the outcroppings. No specimens were observed deep within the caves. Most lizards were observed on vertical surfaces in shaded areas and would retreat into nearby cracks at the slightest provocation. Males would often curl their tail up over their back and wave the bright yellow (regenerated) or white (original) posterior section from side to side. Relationships. Cnemaspis paripari and its sister species of C. nigridia from the nigridia species group (Fig. 2). Material examined. East Malaysia: Sarawak, Gua Pari-pari ZRC 2.6812; Gua Angin LSUHC 9185, ZRC 2.6813–14 (type series).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 101 137 FIGURE 53. Cnemaspis paripari from Gua Angin, Sarawak, East Malaysia. Upper left: adult male (LSUDPC 4924) in the dark color pattern phase. Upper right: gravid female (LSUDPC 4923) in the dark color pattern phase. Lower: karst microhabitat at Gua Angin. Photographs by LLG.

Kendallii group. The kendallii group is a well-supported lineage containing a morphologically diverse group of endemic, insular species from the Seribuat (Cnemaspis baueri and C. pemanggilensis) and Natuna (C. mumpuniae sp. nov. and C. sundainsula sp. nov.) archipelagos along with C. bidongensis from Pulau Bidong from Peninsular Malaysia; C. peninsularis sp. nov. from Peninsular Malaysia and Singapore, and C. kendallii from East Malaysia and Indonesia (Figs. 2, 4). As noted above, the monophyly of this group is further supported in that these seven species lack precloacal pores (Fig. 5). The relationships within the kendallii group clearly indicates that C. kendallii is polyphyletic, being that it has at least six separate, independent origins (Fig. 2 and see below): C. kendallii from Borneo; C. kendallii from Peninsular Malaysia (= C. peninsularis sp. nov.), and C. kendallii from the Natuna Archipelago, Indonesia (Gonatodes kendallii fide De Rooij [1915] = C. mumpuniae sp. nov. and Gonatodes kendallii fide Günther [1895] = C. sundainsula sp. nov.). The precise phylogenetic placement of C. kendallii (Gonatodes kendallii fide Smedley [1928] = C. sundagekko sp. nov.) from Pulau Siantan of the Anambas Archipelago, Indonesia is not yet known but will likely add an additional independent origin (see below). The polyphyletic nature of C. kendallii was first noted by Grismer et al. (2008b) and Grismer (2011a:334). Leong et al. (2003) indicated that populations on a number of Indonesian Islands were also likely to be different species. In their revision of C. kendallii, Das & Bauer (1998) considered the population from the upland region of Bukit Larut in Peninsular Malaysia, all Peninsular Malaysian and Seribuat Archipelago populations, and the Natuna Besar and Anambas Island populations to compose C. kendallii which in fact is a composite of six species (Grismer et al. 2008b and herein), some of which occur in different clades. The phylogeny also indicates that the Peninsular

102 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 138 Malaysian and Seribuat Archipelago populations (i.e. C. peninsularis sp. nov.) and the Natuna Besar Island populations are not closely related to Bornean C. kendallii, which which is the provenance of the holotype (Gray 1845). Additionally, examination of the population from Pulau Siantan from the Anambas Archipelago reveals that it too is not conspecific with Bornean C. kendallii but likely related to a clade containing C. baueri, C. pemanggilensis, C. mumpuniae sp. nov., C. bidongensis, and C. peninsularis sp. nov. being that it has the derived character state of caudal tubercles encircling the tail. Therefore, C. kendallii sensu lato is reclassified and the new species are described below. In the molecular analysis, Cnemaspis sundainsula sp. nov. is the basal species in a well-supported monophyletic group comprised of it and C. pemanggilensis, C. kendallii sensu stricto, C. baueri, C. mumpuniae sp. nov., C. bidongensis, and C. peninsularis sp. nov. The monophyly of these latter species is supported in that they are the only species of Cnemaspis that have caudal tubercles encircling the tail and the monophyly of C. baueri, C. mumpuniae sp. nov., C. bidongensis, and C. peninsularis sp. nov. is supported in that they are the only Cnemaspis in which the posterior portion of the original tail is black. The kendallii group is diagnosed by having a maximum SVL 58.1–84.5 mm; 9–13 supralabials; 7–12 infralabials; keeled ventral scales; 0–6, contiguous, round, pore-bearing, no precloacal pores; body tuberculation moderate; 17–37 paravertebral tubercles; caudal tubercles not restricted to a single paravertebral row; lateral row of caudal tubercles present; 1–4 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; and 25–38 subdigital fourth toe lamellae.

Cnemaspis sundainsula sp. nov. Sunda Island Rock Gecko Fig. 54

Gonatodes kendallii Günther 1895:500; De Rooij 1915:26 (in part) Cnemaspis cf. nigridia Leong, Grismer & Mumpuni 2003:170 Cnemaspis kendallii Das & Bauer 1998:13

Holotype. Adult male MZB. Lace.9438 collected by Awal Riyanto on 24 October 2011 from Mount Ranai, Bunguran Timur district , Natuna Regency , Kepulauan Riau Province, Bunguran (Great Natuna) Island, Indonesia (03o57’24.5”N, 108o21’08.3”E) at 345 m above sea level . Paratypes. Adult male paratypes MZB.Lace 9436–37 and adult female paratypes MZB.Lace 9439–40 have the same data as the holotype except that MZB.Lace 9440 was collected on 25 October 2011; adult maleTNHC 64276, adult female TNHC 62277 and adult male MZB.Lace.4621 were collected on 3 April 2003 by B. J. Evans, Mohd. Iqbal Setiadi and Gandhi Probowo from Mount Ranai, Bunguran Timur District, Natuna Regency , Kepulauan Riau Province, Bunguran Island, Indonesia (03°57.381’ N, 108°21.319’ E); adult female USNM28139 was collected on 2 July1900 by W. L. Abbott from Bunguran (=Pulau Natuna Besar), Kepulauan Riau Province, Indonesia (03°57.381’ N, 108°21.319’ E); adult males MZB.Lace 10156 and 10159 were collected on 26 August 2013 and 27 August 2013, respectively by Awal Riyanto and Zamri at Ceruk Forest Conserve, Selemam Village, Bungaran Timur Laut District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia(03o58’31.6” N, 108o17’52.2” E) at 51 m above sea level; and adult males MZB.Lace 10160–61 were collected on 28 August 2013 by Awal Riyanto and Zamri at Gunung Air Hiu Recreation Area, Ceruk Village, Bungaran Timur Laut District, Natuna Regency, Kepulauan Riau Province, Bunguran Island , Indonesia (03o59’03.7” N, 108o19’06.0”E) at 117 m above sea level. Additional specimens examined. Adult males MZB.Lace 10157–58 collected on 26 August 2013 and 27 August 2013, respectively by Awal Riyanto and Zamri at Ceruk Forest Conserve, Selemam Village, Bungaran Timur Laut District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia; subadult males MZB.Lace 9439–40 and MZB.Lace 10162 were collected on 28 August 2013 by Awal Riyanto and Zamri at Gunung Air Hiu Recreation Area, Ceruk Village, Bungaran Timur Laut District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia. Diagnosis. Maximum SVL 84.5 mm; 8–11 supralabials; 7–10 infralabials; ventral scales keeled; no precloacal pores; 26–37 paravertebral tubercles; tubercles linearly arranged, preent on flanks but absent in lateral caudal

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 103 139 furrows; ventrolateral caudal tubercles present; lateral row of caudal tubercles present; caudal tubercles not encircling tail; subcaudals smooth but bearing an enlarged median row of smooth scales occasionally posteriorly; 2–4 postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of first toe weakly enlarged to enlarged; subtibials keeled; 25–29 subdigital fourth toe lamellae; small, light-colored round sponts on flanks; gukar region, throat, and lateral sections of abdomen orange; anterior subcaudal region yellow, posterior region white (Tables 6,7).

FIGURE 54. Cnemaspis sundainsula sp. nov. from Mount Ranai, Kecamatan Bunguran Timur, Kabupaten Natuna, Propinsi Kepulauan Riau, Bunguran (Great Natuna) Island, Indonesia. Upper left: granite bolder habitat in primary forest at the type locality at the base of Mount Ranai. Upper right: adult male (LSUDPC 8790) in dark color phase. Middle left: adult female (LSUDPC 8796) in situ displaying orange flanks and undersides. Middle right: adult male (LSUDPC 8813) in situ at night in light color phase. Lower left: holotype MZB.Lace.9438 (upper left) and paratypes MZB.Lace 9436, 9437 and 10161 (upper row from left to right) and paratypes MZB.Lace 10156, 10160, 10159, and 4621 (lower row from left to right). Lower middle, upper: paratypes holotype USNM 28136 (left) and paratypes THNC 64276–77 (middle and right, respectively). Lower middle, lower: underside of adult male (uncataloged). Lower right: hatchling (LSUDCP 8829) in situ. Photographs by LLG.

104 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 140 Description of holotype. Adult male, SVL 84.1 mm; head robust, oblong in dorsal profile, moderate in size (HL/SVL 0.23), not narrow (HW/SVL 0.15), flattened (HD/HL 0.41), distinct from neck; snout moderate (ES/HL 0.42), slightly concave in lateral profile; postnasal region constricted medially; scales of rostrum keeled, raised, larger than scales on occiput; distinct, supraorbital ridges; no frontorostral sulcus; canthus rostralis rounded; eye large (ED/HL 0.22); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, posterior 90% divided by longitudinal groove; rostral bordered posteriorly by two large supranasals and external nares, laterally by first supralabials; 9R,L raised supralabials decreasing in size posteriorly; 8R,L infralabials, decreasing in size posteriorly; nostrils round, oriented dorsoposteriorly; mental large, triangular, flat, bordered posteriorly by six postmentals, first two on either side largest; gular and throat scales granular, keeled, raised; pectoral scales slightly larger. Body robust (AG/SVL 0.43); small, granular, rugose, dorsal scales generally equal in size throughout body, intermixed with larger, multicarinate tubercles more or less linearly arranged; tubercles extend from occiput to base of tail; tubercles on flanks; 32 paravertebral tubercles; pectoral and abdominal scales small, granular, keeled, same size throughout; abdominal scales slightly larger than dorsals; no precloacal pores; forelimbs moderately long, robust (FL/SVL 0.19); dorsal scales of brachium raised, keeled; dorsal scales of forearm raised, keeled; ventral scales of brachium keeled, raised, juxtaposed; ventral scales of forearm keeled, raised, juxtaposed; palmar scales, juxtaposed, raised, broadly keeled; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout digit; interdigital webbing absent; fingers increase in length from first to fourth with fourth longer than fifth; hind limbs robust, slightly longer and thicker than forelimbs (TBL/SVL 0.24); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales raised, keeled, juxtaposed, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; weakly enlarged to enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched, wide throughout digit; interdigital webbing weak to absent; toes increase in length from first to fourth with fourth being slightly longer than fifth; 28 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales raised, keeled, juxtaposed; deep middorsal and lateral caudal furrows; subcaudal scales smooth; median row of enlarged subcaudal scales posteriorly; caudal tubercles do not encircle tail; tubercles absent from lateral furrows; three enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail. Coloration in life (Fig. 54). Dorsal ground color of head, body and limbs yellowish brown; paired, yellowish, lineate markings on rostrum; irregularly shaped yellowish markings on top of head; thin, black, upper postorbital stripe extending to occiput; thin, black, lower postorbital stripe extending onto flank; thin, yellowish, postorbital stripe highlighting row of tubercles on latter surface of occiput; transverse, beige marking on nape followed by one square and two rectangular medial beige markings on body alternating with small, elongate, thin, black vertebral markings; light rectangular markings on body grade into diffuse, light caudal bands alternating with darker brown bands; caudal bands do not encircle tail; diffuse light blotches on flanks; tubercles on flanks white; dark, diffuse, rectangular markings on back and flanks alternate with light markings; gular region and throat yellow-orange; lateral margins of abbomen and lower flanks yellow-orange; anterior one-half of subcaudal region yellow, posterior one-half white; ventral surfaces of pectoral region and limbs beige. Variation (Fig. 54). All paratypes closely resemble the holotype in coloration and pattern although TNHC 64276–77 and MZB.Lace 4621 are not nearly as boldly marked. The color pattern of MZB.Lace 10156 is much bolder than that of the holotype in that the dark and light dorsal markings stand in distinct contrast to one another. The dark dorsal markings of MZB.Lace 10161 are elongate as opposed to being more square to roundish as in the other specimens. TNHC 64277 lacks a tail and the tail of TNHC 64276 is regenerated and composed of small, dark, roundish, juxtaposed scales that are weakly keeled on the dorsal surface whereas the subcaudals are smooth beige, and slightly larger. USNM 28139 is badly faded and only a general color pattern that matches that of the holotype is visible. Hatchlings tend to have yellow undersides. Meristic differences are listed in Table 12. Comparisons. Within the Southern Sunda clade, Cnemaspis sundainsula sp. nov. is differentiated from the species of the nigridia group (C. nigridia and C. paripari) by having a greater maximum SVL (84.5 mm versus 50.7–75.5 mm); lacking as opposed to having precloacal pores; having tubercles on the flanks as opposed to lacking them; having smooth as opposed to keeled subcaudals; generally lacking as opposed to having a median row of enlarged, subcaudal scales; and lacking as opposed to having weakly enlarged, metatarsal scales beneath the first toe. From the species of the kendallii group (C. baueri, C. pemanggilensis, C. mumpuniae sp. nov., C.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 105 141 bidongensis, and C. peninsularis sp. nov.) of which it is a member, it can be differentiated by having a much greater maximum SVL (84.5 mm versus 58.1–76.0 mm), having smooth as opposed to keeled subcaudal scales, and not having caudal tubercles that encircle the tail. From C. limi, C. sundainsula sp. nov. is separated by having linearly as opposed to randomly arranged dorsal tubercles, and having a ventrolateral row of caudal tubercles. Cnemaspis sundainsula sp. nov. can be differentiated from C. sundagekko sp. nov, by having a much larger maximum SVL (84.5 mm versus 68.0 mm), a greater number of paravertebral tubercles (26–37 versus 20–25), smooth as opposed to keeled subcaudals, caudal tubercles that do not encircle the tail, and fewer subdigital lamellae on the fourth toe (26–31 versus 33–38).

TABLE 12. Meristic and mensural character states of the type series of Cnemaspis sundainsula sp. nov. w = weak; f = female; m = male; post = posterior; / = data unavailable; and r = regenerated. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right. USNM TNHC TNHC MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace

28139 64276 64277 9438 9436 9437 10161 10156 10160 10159 4621

paratype paratype paratype holotype paratype paratype paratype paratype paratype paratype paratype Supralabials 10 9 10 9 9 9 11 8 10 9 10 Infralabials 8 7 9 8 10 8 9 9 8 8 9 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 1 1 1 1 1 No. of precloacal pores 0 0 0 0 / 0 / 0 0 0 / Precloacal pores continuous (1) or separated (0) / / / / / / / / / / / Precloacal pores elongate (1) or round (0) / / / / / / / / / / / No. of paravertebral tubercles 35 31 37 32 31 34 26 33 31 29 31 Tubercles linearly arranged (1) or more random (0) 1 1 1 1 1 1 1 1 1 1 1 Tubercles present (1) or absent (0) on flanks 1 1 1 1 1 1 1 1 1 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 / / 0 0 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 / / 1 1 1 1 1 1 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 / / 1 1 1 1 1 1 1 1 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 / / 0 0 0 0 0 0 0 0 Subcaudals keeled (1) or not (0) 0 / / 0 0 0 0 0 0 0 0 Single median row of keeled subcaudals (1) or not (0) 0 / / 0 0 0 0 0 0 0 0 Caudal tubercles encircle tail (1) or not (0) 0 / / 0 0 0 0 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 / / w,post w,post w,post / w,post post / w,post Postcloacal spurs 4 4 2 3 / / 3 3 2 2 / Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 0 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 1 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) w w w 1 1 1 1 1 1 1 1 No. of 4th toe lamellae 28 28 29 25 26 27 28 26 27 26 29 Sex f m f m f f f m m m f SVL 80 81 79.7 84.5 77.5 82.3 75.2 75.2 77.7 75.2 68.9 TL 97.6 65.9 / 82.4/12.8 75.6/14.8 69.3/19.3 34.6/35.1 93.8 90.4 38.9/42.3 83.5 TW 8.9 9.6 10.6 8.9 9.8 8.5 7.9 7.8 8.4 7.5 6.9 FL 15.4 16.8 15.7 15.7 16.9 15.1 14.7 13.8 14.6 14.8 13.9 TBL 19.1 19.2 19.1 20.7 20.9 19.6 17.5 18.5 18.8 18.2 16.6 AG 34.5 34.2 30.8 35.9 33.4 33.1 32.1 32.7 31.9 31.7 27.9 HL 18.4 20 20.3 20.6 20.4 20.6 19.2 20.1 19.8 20.3 17.4 HW 13.6 13.4 14.6 13.9 12.9 12.6 12.7 12.8 12.8 12.9 12.1 HD 8.9 8.9 9.2 9.1 8.5 7.8 8.1 8.2 8.1 7.1 7.5 ED 4.2 3.9 4.2 4.5 4.3 4.3 3.9 3.6 3.9 3.7 3.5 EE 6.1 2.5 2.3 6.3 4.9 6.1 5.7 5.9 5.6 5.9 5.3 ES 10 9.7 9.6 10.1 9.6 10.1 9.3 9.5 9.1 8.2 8.4 EN 7.7 7.5 7.4 7.6 7.2 7.9 6.9 7.4 6.9 6.3 6.6 IO 1.9 4 4.5 4.9 4.5 4.8 4.5 4.7 4.5 4.7 4.5 EL 1.7 2.4 2.2 2.2 2.1 2.4 1.9 1.7 2.1 1.2 2.1 IN 1.8 1.7 1.8 2.1 1.7 1.9 1.7 1.8 1.9 1.8 1.9 Ó

Distribution.Cnemaspis sundainsula sp. nov. is known only from Bunguran Island (= Pulau Natuna Besar/ Great Natuna) of the Natuna Archipelago, Kepulauan Riau Province, Indonesia (Fig. 4) but is likely to occur on nearby islands in the archipelago. Natural History. Cnemaspis sundainsula sp. nov. inhabits primary and secondary forests along the base of

106 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 142 Mount Ranai from at least 51–345 m above sea level (De Rooij 1915; Günther 1895). It is not certain if this species extends up to the summit but if granite boulders are present we suspect it does. Lizards occur almost exclusively on large granite boulders (Fig. 54) and are only rarely found on tree trunks. This species is abundant and active during the day while in its dark color phase (Fig. 54) and often in male-female pairs. While active during the day, lizards remain wary and will not venture too far out onto the exposed boulder surfaces but rather remain near cracks or narrow spaces between adjacent boulders were escape is possible. Lizards are commonly seen upside down suspended from the undersides of boulders with their limbs outstretched displaying their birght-orange lower flanks, throat, and abdomens (Fig. 54). Before escaping into cover lizards, roll their tails up over their backs and display the immaculate, white posterior subcaudal region (Fig. 54). In dark crevices and boulder spaces, often the subcaudal region is all that can be seen. At night, lizards venture farther out onto the exposed boulder surface and sequester themselves in a shallow depression or crease or along the edge of a small ridge where they are generally inactive. During this period, lizards are in their light color phase (Fig. 54) and are much more approachable. We have observed gravid females carrying two eggs, hatchings, and juvenlies during April. Cnemaspis sundainsula sp. nov. occurs syntopically with C. mumpuniae sp. nov. and Cyrtodactylus hikidai Riyanto when the latter two species occur on granite boulders. The ecological pattern of temporal partitioning on islands seen between C. sundainsula sp. nov. and Cyrtodactylus hikidai is similar to that between C. psychedelica on Hon Khoai Island, Vietnam which is diurnal and found syntopically with the nocturnal Cyrtodactylus sp. nov. 1 (Grismer et al. 2010b); C. boulengerii on Con Son Island in the Con Dao Archipelago, Vietnam which is diurnal and syntopic with the larger Cyrtodactylus condorensis (Smith); and the large, diurnal, granite boulder-dwelling C. limi from Pulau Tioman, Peninsular Malaysia which is syntopic with the larger nocturnal Cyrtodactylus tiomanensis Das & Lim. Additionally, C. sundainsula sp. nov. and C. mumpuniae sp. nov. partition their habitat by SVL, elevation, and general microhabitat preference where C. sundainsula sp. nov. is a large, rock-dweller and C. mumpuniae is the smaller, habitat generalist. A parallel system exists on Pulau Tioman, Malaysia with the smaller, habitat generalist C. peninsularis sp. nov. and the larger, rock-dwelling C. limi (Grismer 2011a). Etymology. The specific epithet sundainsula is derived from the word Sunda which originally referred to a Hindu Kingdom in western Java existing from 669–1579. Sunda is now commonly used as an adjective associated with particular geographic features in the western regions of Southeast Asia associated with the South China Sea and its fringing continental areas. The Latin insula (singular) means island and sundainsula is an invariable noun in apposition in reference to this species being endemic to an island on the submerged Sunda Plains. Relationships. Cnemaspis sundainsula sp. nov. is part of the kendallii group within the Southern Sunda clade and is basal to a group containing C. pemanggilensis, C. kendallii sensu stricto, C. baueri, C. mumpuniae sp. nov., C. bidongensis, and C. peninsularis sp. nov. (Fig. 2).

Cnemaspis kendallii (Gray, 1845) Kendall’s Rock Gecko Figs. 55, 56

Heteronota kendallii Gray, 1845:174 (in part) Gonatodes kendalli (in part) Boulenger, 1885:63; Shelford, 1901:48 Gonatodes affinis Shelford, 1901:49 Gonatodes kendallii de Rooij, 1915:25 (in part)

Lectotype. BM XXII.92 (designated by Dring 1979:223). Type locality: “Borneo” (02°34.44 N, 104°19.53 E) at 100 m in elevation. Diagnosis. Maximum SVL 58.4 mm; 10 or 11 supralabials; eight or nine infralabials; keeled ventral scales; no precloacal pores; 18–26 paravertebral tubercles; body tubercles semi-linearly arranged, weak to present on flanks, tubercles absent from lateral caudal furrows; ventrolateral and lateral row of caudal tubercles present; caudal tubercles encircle tail; subcaudals keeled with no enlarged median row; two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials keeled; 25–33 subdigital fourth toe lamellae; distinct black and white caudal bands on posterior portion of tail; subcaudal region immaculate white (Tables 6,7).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 107 143 FIGURE 55. Cnemaspis kendallii. Left: adult female (LSUDPC 4903) from Santubong, Sarawak, East Malaysia in the light color pattern phase. Upper right: adult male (LSUDPC 4875) from Gunung Gading, Sarawak, East Malaysia in the dark color pattern phase. Photographs by LLG. Lower right: adult male (LSUDPC 6804) from Pulau Karimata, Kalimantan, Indonesia. Photograph by Umilaela Arafin.

Color pattern in life (Figs. 55, 56). Coloration differs greatly depending on the time of the day. Diurnal coloration: dorsal ground color of the head, body, and limbs grey to dark brown; dark and yellow markings on top of head; thin, dark, diffuse, postorbital stripe extends onto nape; medial, black marking on nape followed by black, vertebral spots extending from shoulder region to base of tail and flanked laterally by additional row of spots on each side on anterior portion of body; dorsum and upper portions of limbs bearing whitish to yellowish spots, those in vertebral region largest; black and white caudal bands in males and females, bands encircling tail in females; subcaudal region nearly immaculate white in males in both original and regenerated tails; dorsal pattern of regenerated tail beige with dark flecking; ventral surfaces of head and neck dull-beige; pectoral region, abdomen and ventral surfaces of limbs beige, usually immaculate. Nocturnal coloration: ground color of dorsal surface of head and body nearly white to light yellow, highlighting dark markings on head and spotting on dorsum; ground color of limbs and tail yellowish. Distribution. Cnemaspis kendallii ranges throughout northwestern Borneo in Sarawak, East Malaysia (Das & Bauer 1998) and western Kalimantan, Indonesia. It ranges northward to Pulau Serasan of the Southern Natuna Islands and onto Pulau Buona of the Tambelan Islands, Pulau Pedjantan [sic.] (=Pejantan), and Pulau Karimata (Umilaela et al. 2009) to the south (Fig. 4). Das and Bauer (1998) erroneously reported this species from “Pulo [sic] Lingga” based on specimen USNM 28145. However, Pulau Lingga is an island south of Pulau Bintan, Indonesia that lies just south of Singapore on the western edge of the South China Sea. The original hand written collection label on USNM 28145 reads “Pulau Lingung [=Pulau Lagong] near Natuna Besar” which is a small island off the southern tip of Pulau Natuna Besar (Fig. 4). USNM 28145 is recognized here as C. mumpuniae sp. nov.

108 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 144 FIGURE 56. Upper: adult male Cnemaspis kendallii (LSUDPC 4901) at Santubong, Sarawak, East Malaysia in the dark color pattern phase displaying the white underside of the original tail. Lower: habitat of Cnemaspis kendallii at Matang, Sarawak, East Malaysia. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 109 145 FIGURE 57. Upper: adult male Cnemaspis pemanggilensis (LSUDPC 2578) from Pulau Pemanggil, Johor, Peninsular Malaysia in the dark color pattern phase. Lower: habitat on Pulau Pemanggil. Photographs by LLG.

110 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 146 Natural history. Cnemaspis kendallii is a habitat generalist that ranges throughout primary, secondary, and old secondary forests. Lizards are generally diurnal and can be found on the shaded surfaces of large granite boulders, limestone formations, tree roots, and tree trunks (Fig. 56). Upon retreat, males often roll their tail over their back displaying its white, immaculate underside (Fig. 56) while waving the tip back and forth. At Gunung Gading, we observed lizards during the day on the same granite boulders wherein we observed the much larger C. nigridia at night, indicating these species may be partitioning their microhabitat by means of body size and activity period as has been suggested for other sympatric pairs of Cnemaspis (Grismer et al. 2010b). At night, C. kendallii is often found abroad sleeping on tree trunks and other vegetation and in open areas on the faces of granite boulders. Relationships. Cnemaspis kendallii is the sister species of C. pemanggilensis (Fig. 2). Remarks. We examined two specimens from the Tambelan Archipelago, Indonesia collected in 1899: one from Pulau Benua (=Buona) and another from Pulau Pejantan (USNM 26573 and 26555, respectively). Both were very faded and in poor shape overall. More importantly, both lacked tails meaning that eight of some of the most important diagnostic character states used to delimit species boundaries between Cnemaspis were unavailable for examination. The combination of the remaining character states such as no precloacal pores, keeled ventrals and subtibials, SVLs of 51.5 and 55.7 mm coupled with their locality off the west coast of Borneo would place them in C. kendallii as was reported by Das and Bauer (1998). However, being that this archipelago has been separated from Borneo for nearly as long as the Anambas and Natuna archipelagos (which collectively harbor at least three species of endemic Cnemaspis) and their specific identity is not possible, we recognize these populations as C. cf. kendallii. Plans are in preparation to collect additional specimens. Material examined. East Malaysia: Sarawak, Gunung Gading LSUHC 9171–73, 9176, USNM 76633; Santubong LSUHC 9178–81. Indonesia: Riau Province, Natuna Archipelago, Pulau Serasan TNHC 64278; Tambelan Archipelago, Pulau Buona USNM 26573; Pulau Pejantan USNM 26555.

Cnemaspis pemanggilensis Grismer & Das, 2006 Pemanggil Island Rock Gecko Fig. 57

Holotype. ZRC 2.6043. Type locality: “Batu Buau, a small rocky hill behind Kampung Buau on the west side of Pulau Pemanggil, Johor, West Malaysia” (02°34.44 N, 104°19.53 E) at 100 m in elevation. Diagnosis. Maximum SVL 76.0 mm; 10–13 supralabials; 8–10 infralabials; keeled ventral scales; no precloacal pores; 30–37 paravertebral tubercles; body tubercles randomly arranged, weak to absent on flanks, absent from lateral caudal furrows; ventrolateral row of caudal tubercles present anteriorly; lateral row of caudal tubercles present; caudal tubercles encircle tail; subcaudals keeled, bearing an enlarged median row of keeled scales; one or two postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; distal submetatarsal scales of fourth toe enlarged; subtibials keeled; 27–34 subdigital fourth toe lamellae; small, yellow spots on flanks (Tables 6,7). Color pattern in life (Fig. 57). Ground color of head, body, limbs, and tail grey; dark, bifurcating, medial stripe on snout; single, dark, preorbital stripe; three dark, postorbital stripes with dorsalmost extending onto nape and forming a tripartite band; middle stripe extends onto nape forming a second tripartite band posterior to and larger than the first; ventralmost stripe extends onto upper portions of forelimb insertions; dark, transversely arranged spots extend from nape to base of tail; distinctive, yellowish spots occur on flanks invading lateral portions of abdominal region; weak, banding pattern on limbs; dark, diffuse banding on anterior portion of tail; ventral surfaces beige bearing dark and light spots. During the day, the dorsal ground color is nearly solid black with virtually no discernable pattern. At night, the dorsal ground color changes to light grey which greatly accentuates a prominent series of dark, transverse, body bands and head stripes. Distribution. Cnemaspis pemanggilensis is known only from the island of Pemanggil (Grismer & Das 2006; Grismer et al. 2006; Grismer 2011b; Fig. 4). Natural history. The habitat on Pulau Pemanggil is severely degraded and has no discernable primary forest (Youmans et al. 2002). Nonetheless, the cave-like systems formed by the piling up of large, granite boulders (Fig. 57) on top of one another maintain the microhabitats and refugia necessary to support a dense population of Cnemaspis pemanggilensis (Grismer & Das 2006). According to Grismer (2011a), lizards are found in nearly every

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 111 147 cave system at all elevations from sea level to 250 meters. During the day, lizards remain inside the caves on all vertical and inverted surfaces where they are extremely wary and rapidly move into deep crevices and cracks at the slightest provocation. At night, however, lizards will venture out along the cave openings and on large boulders immediately outside the caves and become much more approachable. Relationships. Cnemaspis pemanggilensis is the sister species to C. kendallii (Figs. 2,5). Material examined. West Malaysia: Johor, Pulau Pemanggil ZRC 2.6043–51 (type series). Material examined since Grismer & Das (2006): West Malaysia: Johor, Pulau Pemanggil LSUHC 4458, 4460, 4464, 4476, 4495, 8011–16.

FIGURE 58. Cnemaspis baueri from Pulau Aur, Johor, Peninsular Malaysia. Upper left: granite boulder microhabitat on Pulau Aur. Upper right: adult female (LSUDPC 958) in the dark color pattern phase. Middle right: juvenile male (LSUDPC 959) in the dark color pattern phase. Lower right: adult male (LSUDPC 961) in the dark color pattern phase. Lower left: adult male (LSUDPC 957) in the dark color pattern phase. Photographs by LLG.

Cnemaspis baueri Das & Grismer, 2003 Bauer’s Rock Gecko Fig. 58

Holotype. ZRC 2.5291. Type locality: “Kampung Berhala (2°27.507 N, 104°30.163 E), Pulau Aur, Johor, West Malaysia” at 50 m in elevation.

112 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 148 Diagnosis. Maximum SVL 67.4 mm; 11–13 supralabials; 8–12 infralabials; keeled ventral scales; no precloacal pores; 18–27 paravertebral tubercles; body tubercles randomly arranged, absent to weakly present on flanks; tubercles absent from lateral caudal furrows; ventrolateral and lateral caudal rows of tubercles present; caudal tubercles encircle tail; subcaudals keeled; a median row of enlarged, keeled subcaudals present; one or two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials keeled; 26–32 subdigital fourth toe lamellae; uniform brown dorsal ground color; thin, yellow reticulum on occiput and nape; large, black, round spots on nape and anterior portion of body; thin, yellow, caudal bands anteriorly; posterior portion of original tail black in males; anterior caudal tubercles white (Tables 6,7). Color pattern in life (Fig. 58). Ground color of dorsal surface of head, body, and limbs dark-brown to olive; thin, yellow reticulum on the top of head; dark, postorbital stripe extends onto nape; medial, dark marking on nape; black, shoulder patches absent; 5–7 dark, vertebral blotches flanked in shoulder region by elongate, dark blotches followed by dark spots extending onto midsection of body; limbs and body faintly mottled with slightly lighter coloration; posterior two-thirds of tail black; anterior one-third encircled by large, cream-colored tubercles; regenerated tail unicolor brown; ventral surfaces of head, body, limbs, and anterior one-third of tail dull beige, immaculate; sexual dimorphism absent. Distribution. Cnemaspis baueri is known only from Pulau Aur, Johor and the nearby rocky island of Dayang (Grismer 2011a; Fig. 4). Das (2010) erroneously reports this species as being endemic to Pulau Tulai, Johor. Natural History. Grismer (2011a) reported that Cnemaspis baueri is a saxicolous gecko common on rocky outcroppings in primary and secondary, lowland, coastal forests where it is found almost exclusively on the vertical surfaces of large granite boulders, within deep crevices, and within cave-like microhabitats formed by the aggregation of large boulders (Fig. 58). During the day, the activity of C. baueri is restricted to the shaded surfaces of large boulders under the forest canopy or within the cave-like environments wherein lizards can be found in surprisingly high densities. At night, lizards venture farther out into the open onto all surfaces of the rocks but are far less active. Das & Grismer (2003) estimated finding 200–250 egg scars on the underside of a large boulder in a communal laying site. Females carrying two eggs have been observed during July (Grismer 2011a). Relationships. Cnemaspis baueri is the sister species of a monophyletic group containing C. mumpuniae sp. nov., C. bidongensis, and C. peninsularis sp. nov. (Figs. 2, 5). Material examined. West Malaysia: Johor, Pulau Aur ZRC 2.5291–99 (type series). Material examined since Das and Grismer (2003): West Malaysia: Johor, Pulau Aur LSUHC 3921–24, 4700–01, 4717–23, 4725, 4727, 4729, 4744, 4808, 7272–74, 7301–03, 7319, ZRC 2.5093, 2.5095–96.

Cnemaspis mumpuniae sp. nov. Mumpuni Rock Gecko Fig. 59

Cnemaspis kendallii Das & Bauer 1998:12 (in part) Cnemaspis cf. kendalli Leong, Grismer & Mumpuni 2003:170

Holotype. Adult male MZB.Lace 10167 collected by Awal Riyanto and Zamri on 31 August 2013 from Sekunyam Forest Reserve, Mekarjaya Village, Bunguran Barat district, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia (03°40’30.8”N; 108°09’18.2”E) at 80 m above sea level. Paratypes. Adult male MZB.Lace 10166 bears the same data as the holotype. Adult male MZB.Lace 10163 collected by Awal Riyanto and Zamri on 3 September 2013 respectively from Harapan Jaya Village, Bunguran Tengah District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia (03°51’29.8”N; 108°17’59.3”E) at 46 m above sea level, adult maleMZB.Lace.9441collected by some collectors of holotype 24 October 2011, from Mount Ranai, Bunguran Timur District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia (03o57’24.5”N, 108o21’08.3”E) at 345 m above sea level. Adult male MZB.Lace 10169 and adult female MZB.Lace 10168 collected by Awal Riyanto and Zamri on 1 September 2013 from Teluk Lampa Forest, Pulau Tiga District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia (03°40’04.9” N; 108°08’20.2” E) at 10 m above sea level.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 113 149 FIGURE 59. Cnemaspis mumpuniae sp. nov. from Pulau Natuna Besar. Upper left: Adult male (LSUDPC 8784) from Gunung Ranai. Upper middle: in situ adult female (LSUDPC 8815) from Gunung Ranai. Upper right: adult male (LSUDPC 8784) from Selamam. Middle left: hatchling (LSUDPC 8823) from Gunung Ranai. Lower left: in situ adult male (LSUDPC 8784) from Gunung Ranai. Lower right: type series from Sekunyam Forest Reserve, Desa Mekarjaya, Kecamatan Bunguran Barat, Kabupaten Natuna, Kepulauan Riau Province, Indonesia, MZB.Lace 10167 (holotype) far left and paratypes 10166, 9941, 10169, 10168, and 10163 from left to right. Photographs by LLG.

Additional specimens examined. Adult female MZB.Lace 10155 collected by Awal Riyanto and Zamri on 25August 2013 from a fragmented forest area at Bedung Village, Bunguran Tengah District, Natuna Regency, Kepulauan Riau Province, Bunguran Island, Indonesia (03°56’36.8” N; 108°13’19.9” E) at 41 m above sea level. Adult females MZB.Lace 10164–65 have the same data as the holotype. Diagnosis. Cnemaspis mumpuniae sp. nov. differs from all other species of Cnemaspis in having a maximum SVL reaching 56.6 mm SVL; 10 or 11 supralabials; 8–11 infralabials; keeled ventrals; no precloacal pores; moderate dorsal tuberculation; 18–24 paravertebral tubercles; dorsal body tubercles semi-linearly arranged; weak tuberculation on flanks; caudal tubercles encircling tail; tubercles absent from lateral caudal furrows; ventrolateral and lateral row of caudal tubercles present; subcaudals keeled; single, median row of enlarged subcaudals; one or two postcloacal tubercles on either side of base of tail; no enlarged femoral, subtibial or submetatarsal scales; subtibials usually keeled; 29–35 subdigital lamellae on fourth toe; thin, white, nuchal loop; dorsal ground color brick-red; small, light, round spots on flanks; regenerated tail yellow; posterior portion of original tail black in males (Tables 6,7). Description of holotype. Adult male SVL 51.6 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.26), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.41), distinct from neck; snout short (ES/HL 0.50), concave in lateral profile; postnasal region weakly constricted medially, flat; scales of rostrum weaklykeeled, slightly raised, slightly larger than similarly shaped scales on occiput; low, supraorbital ridges; weak frontorostral

114 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 150 sulcus; canthus rostralis not very discernable; eye large (ED/HL 0.23); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 90% divided by longitudinal groove; rostral bordered posteriorly by two large supranasals, an equally sized azygous scale, and external nares; boredered laterally by first supralabials; 10R,L raised supralabials decreasing in size posteriorly; 8R,L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, concave medially, bordered posteriorly by two large, rectangular, lateral postmentals of similar size and one smaller azygous scale; gular scales raised, smooth; throat scales larger, raised, weakly keeled. Body slender, elongate (AG/SVL 0.43); small, keeled, dorsal scales generally equal in size throughout body, intermixed with larger, multicarinate tubercles in semi-linearly arranged; tubercles extend from occiput to base of tail; tubercles moderate in size; tuberculation weak on lower flanks, 18 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales slightly larger than dorsals; no precloacal pores; forelimbs moderately long, slender (FL/SVL 0.21); dorsal scales of brachium raised, keeled; dorsal scales of forearm raised, keeled; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm weakly keeled, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout digit; interdigital webbing present; fingers increase in length from first to fourth with fourth longer than fifth; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.23); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales raised, keeled, juxtaposed, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally but wider distally throughout digit; interdigital webbing present; toes increase in length from first to fourth with fourth being slightly longer than fifth; 32 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales flat anteriorly, keeled, juxtaposed; weak middorsal and deep lateral caudal furrows; subcaudal scales keeled; median row of enlarged, keeled subcaudal scales; caudal tubercles encircle tail; tubercles absent from lateral furrows; two enlarged postcloacal tubercles on lateral surface of hemipenal swellings at base of tail. Color pattern (Fig. 59). Dorsal ground color brick-red; medial, whitish spot on rostrum, canthus rostralis bearing whitish line; thin, white nuchal loop extending from posterior margin of one orbit to the other; thin, white postorbital line below nuchal loop extending obliquely to corner of mouth; paired whitish markings on occiput; small, white, linearly arranged spots on side of neck and nape; large, faint, dark, linearly arranged blotches on anterior portion of body and nape; flanks bearing small, round, white spots thatextend onto lateral margins of abdomen; five whitish bands consisting of a row of three, transversely aligned blotches occur between limb insertions and extend onto anterior one-half of tail transforming into light, caudal bands that alternate with dark bands, posterior one-half of tail black; dorsal surfaces of limbs mottled with white; ventral surface of gular, pectoral, abdominal, and anterior subcaudal region beige; throat and limbs darker; posterior one-half of subcaudal region black; all other ventral surfaces suffused with black stippling in scales. Variation (Fig. 59). The type series shows a modest array of color pattern variation. MZB.Lace 10168 closely resembles the holotype in overall coloration whereas the dorsal ground color in MZB.Lace 10166 and 10168 is lighter and the overall blotching lighter, giving them a less contrasted and spotted appearance. MZB.Lace 10168 is a female and lacks the black posterior caudal region. The posterior one-half of the tail in MZB.Lace 10169 is regenerated and unicolor tan. MZB.Lace 10163 is very faded overall but most likely matches MZB.Lace 9441 and MZB.Lace 10168 in general coloration. The flanks of all specimens of the type series are not as boldly marked as in the holotype. Meristic and mensural variation is listed in Table 13. Comparisons. Cnemaspis mumpuniae sp. nov. is a member of the Southern Sunda clade which includes C. limi, C. nigridia, C. paripari, C. kendallii, C. sundainsula sp. nov., C. pemanggilensis, C. baueri, C. bidongensis, and C. peninsularis sp. nov. Within this clade, it is part of an unresolved polytomy that includes C. kendallii, C. sundainsula sp. nov., C. pemanggilensis, C. baueri, C. bidongensis, and C. peninsularis sp. nov. of the kendallii group (Fig. 2). Cnemaspis mumpuniae sp. nov. is easily separated from C. limi by being much smaller (maximum SVL 56.6 mm versus 88.2 mm); having fewer paravertebral tubercles (18–24 versus 25–35); having keeled versus smooth subcaudal scales; the presence versus the absence of a ventrolateral row of caudal tubercles; having caudal tubercles that encircle the tail versus not having tubercles encircling the tail; and lacking versus having white caudal tubercles. From C. paripari, C. mumpuniae sp. nov. lacks precloacal pores as opposed to having them; has

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 115 151 fewer paravertebral tubercles (18–24 versus 26–31); has as opposed to lacks tubercles on the flanks; has as opposedto lacks a ventrolateral row of caudal tubercles; has caudal tubercles that encircle the tail versus not having tubercles encircling the tail; lacks as opposed to having an enlarged, median subcaudal scale row; and males lack as opposed to having a yellow head, limbs, and back and the posterior one-half of the original tail being white. Within the kendallii group, C. mumpuniae sp. nov. is distinguished from C. sundainsula sp. nov., C. pemanggilensis, and C. baueri by being much smaller (maximum SVL 56.6 mm versus 67.4–84.5 mm) and from C. sundainsula sp. nov. it is further separated by having caudal tubercles that encircle the tail rather than not having such tubercles. Cnemaspis mumpuniae sp. nov. is further separated from C. pemanggilensis by having fewer paravertebral tubercles (18–24 versus 30–37) and lacking as opposed to having an enlarged, median row of keeled subcaudal scales. From C. baueri, C. mumpuniae sp. nov is further differentiated by lacking an enlarged, median row of keeled subcaudal scales and not having a uniform brown dorsal color pattern bearing large, elongate black blotches on the nape and shoulder region. Cnemaspis mumpuniae sp. nov. is differentiated from C. kendallii sensu stricto in that the posterior two-thirds of the original tail in adult male C. mumpuniae sp. nov. is black dorsally and ventrally and in adult male C. kendallii the tail is banded dorsally throughout its length and the subcaudal region is essentially immaculate white. The regenerated tail in adult male C. mumpuniae sp. nov. is yellow and immaculate dorsally and ventrally whereas that of C. kendallii sensu stricto is straw-colored with small black flecks dorsally and the subcaudal region is immaculate white. Additionally, C. kendallii sensu stricto has a row of nearly contiguous tubercles on the lateral margins of the occipital region bordering the nape which are nearly always absent in C. mumpuniae sp. nov. Within the kendallii group, C. mumpuniae sp. nov. is most closely related to the sister species C. bidongensis and C. peninsularis sp. nov. (Fig. 2). It differs from them in having a brick-red ground color and a thin, white, nuchal loop. It is differentiated further from C. peninsularis sp. nov. having as opposed to lacking an enlarged, median, subcaudal scale row.

TABLE 13. Meristic and mensural character states of the type series of Cnemaspis mumpuniae sp. nov. w = weak; f = female; m = male. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right.

MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace

10167 9944 10169 10168 10166 9941

holotype paratype paratype paratype paratype paratype Supralabials 10 10 10 11 11 11 Infralabials 8 10 8 9 10 11 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 No. of precloacal pores 0 0 0 0 0 0 Precloacal pores continuous (1) or separated (0) / / / / / 0 Precloacal pores elongate (1) or round (0) / / / / / 0 No. of paravertebral tubercles 18 19 21 19 19 24 Tubercles linearly arranged (1) or more random (0) 0 w w w w w Tubercles present (1) or absent (0) on flanks w w w w w / Caudal tubercles in lateral furrow (1) or not (0) 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 1 1 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 1 0 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 0 0 0 0 0 Subcaudals keeled (1) or not (0) 1 1 1 1 1 1 Single median row of keeled subcaudals (1) or not (0) 0 0 0 0 0 1 Caudal tubercles encircle tail (1) or not (0) 1 1 1 1 1 1 Enlarged median subcaudal scale row (1) or not (0) 1 1 1 1 1 1 ...... continued on the next page

116 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 152 TABLE 13. (Continued) MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace MZB.Lace

10167 9944 10169 10168 10166 9941

holotype paratype paratype paratype paratype paratype Postcloacal spurs 2 2 2 / 1 0 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 2 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 1 No. of 4th toe lamellae 32 29 30 29 35 30 sex m m m f m m SVL 51.6 56.6 52.5 55 53.6 53.3 TL 58.3 42.1/12.8 14.1/28.8 33.3/22.2 67.5 59.1 TW 6.2 5.6 5.2 5.2 4.9 4.4 FL 10.8 9.9 10.7 10.4 10.3 10.1 TBL 12.1 13.2 13.1 12.2 12.2 13.7 AG 22.1 2.6 23.3 25.4 22.9 24.8 HL 13.7 15.6 13.9 14.2 14.4 16.4 HW 8.7 8.9 8.8 8.9 8.4 9.4 HD 5.6 5.9 6.1 5.7 5.4 6.1 ED 3.2 3.2 3.1 2.8 2.8 2.6 EE 3.7 4.1 4.5 4.2 3.9 3.6 ES 6.8 7.9 6.7 6.6 6.7 6.9 EN 6.1 6.2 5.4 5.4 5.1 5.1 IO 2.9 2.7 3.4 3.2 3.2 4 EL 1 1.1 1.2 1 1.3 0.8 IN 1.4 1.7 1.7 1.5 1.4 1.5

Distribution. Cnemaspis mumpuniae sp. nov. is endemic to the northern group of islands of the Natuna Archipelago, Riau Province, Indonesia. It is known to occur on the islands of Natuan Besar and Lagong but is likely present on many of the other islands as well (Fig. 4). Natural History. Cnemapsis mumpuniae sp. nov. is a diurnal, habitat generalist found in disturbed and undisturbed forests and is widespread throughout Pulau Natuna Besar from sea level to 345 m along the base of Mount Ranai. Lizards occur on both granite boulders and vegetation and are quite adept at substrate matching (Fig. 59). During the day on granite boulders, their ground color is dark-red. On lighter substrates, such as tree trunks in rubber plantations, the ground color can be grayish. At night, when inactive, lizards are nearly white. This species is quite agile and wary during the day, jumping from rock to rock or from trees to rocks to seek shelter in dark crevices and rock spaces. While fleeing, lizards usually elevate the black posterior portion of their tail up over their back and wave it from side to side. At night lizards are quite approachable and can be seen sleeping on the open surfaces of boulders and tree trunks. We observed one lizard sleeping on a leaf at least 5 m above the forest floor. Hatchlings were observed during April. Etymology. The specific epithet recognizes Mrs. Mumpuni, one of the senior herpetologist at the MZB and honors her many contributions over the years to Indonesian herpetology. Relationships. Within the kendallii group, C. mumpuniae sp. nov. is most closely related to the sister species C. bidongensis and C. peninsularis sp. nov. (Fig. 2).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 117 153 Cnemaspis peninsularis sp. nov. Peninsular Rock Gecko Figs. 60, 61

Gonatodes kendalli Flower 1896:833 (in part), 1899:627 (in part); Ridley, 1899:193; Boulenger 1912:38 (in part); Sworder 1925:63; Smith 1930:16 (in part); Nicholls 1949:48 Gonatodes kendallii Smith 1925:23 (in part) Cnemaspis kendalli Henrickson, 1966:55; Bullock 1966:94; Dring, 1979:220; Denzer & Manthey, 1991:313; Lim & Lim, 1992:122 Cnemaspis kendallii Manthey & Grossmann 1997:212 (in part); Das & Bauer 1998:12 (in part); Grandison, 1972:80; Werner & Chou, 2002:185; Das & Grismer, 2003:549; Grismer & Das 2006:5 (in part); Grismer, Youmans, Wood & Grismer, 2006:112; Grismer & Ngo 2007:486 (in part); Baker & Lim, 2008:78; Chan & Grismer 2008:55 (in part); Grismer 2008:30; Grismer & Chan 2008:5 (in part); Grismer, Chan, Nurolhuda & Sumontha 2008a:57 (in part); Grismer, Grismer, Wood & Chan 2008b:24 (in part); Grismer & Chan 2009:30 (in part); Grismer, Norhayati, Chan, Belabut, Muin, Wood, & Grismer 2009:59 (in part); J. Grismer, Grismer & Thou 2010:30 (in part); Grismer, 2010:59 (in part); Grismer & Chan 2010:61 (in part); Grismer, Chan, Quah, Muin, Savage, Grismer, Norhayati, Greer, Remegio 2010c:64 (in part); Grismer, Ngo & Grismer 2010b:57 (in part); Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya 2010a:12 (in part); Grismer 2011a:330 (in part), 2011b:112 (in part); Wood, Quah, Shahrul, & Muin 2013:546 (in part); Grismer, Wood, Amirrundin, Sumarli, Vazquez, Chan, Ismail, Nance, Muhammad, Mohamad, Syed, Kuss, Murdoch & Cobos 2014:449.

FIGURE 60. Cnemaspis peninsularis sp. nov. from Gunung Ledang, Johor, Peninsular Malaysia. Upper right: adult male (LSUDPC 6704) in the dark color pattern phase. Photograph by ESHQ. Lower right: adult male (LSUDPC 4428) in the light color pattern phase. Photograph by LLG. Left: karst microhabitat at Gua Senyum, Pahang, Peninsular Malaysia. Photograph by LLG.

Holotype. Adult female LSUHC 8965 collected on 7 June 2008 by L. L. Grismer, P. L. Wood, Jr., J. L. Grismer and Chan K. O. at 1030 hrs from the base of Gunung Ledang Johor, Peninsular Malaysia (02°20.25’ N, 102°37.11’ E) at 100 m in elevation.

118 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 154 Paratypes. Adult male LSUHC 8966 has the same data as the holotype. Adult male LSUHC 4756 collected on 23 July 2002 by J. L. Grismer at the waterfall on Pulau Tinggi, Johor, Peninsular Malaysia (02°18.013’ N, 104°06.261 E) at 210 m in elevation. Adult male LSUHC 5731 collected on 30 August 2003 by T. A. Youmans on Pulau Babi Besar, Johor, Peninsular Malaysia (02°26.166 N, 103°58.466 E) at 55 m in elevation. Adult male LSUHC 6213 collected on 30 June 2004 by L. Lee Grismer on the Tekek-Juara Trail, Pulau Tioman, Pahang, Peninsular Malaysia (02°48.433 N, 104°9.525 E) at 260 m in elevation. Adult female LSUHC 8210 and adult male LSUHC 8126 collected on 4 September 2006 by L. L. Grismer at Selai, Lubuk Tapah, Endau-Rompin, Johor, Peninsular Malaysia (02°25.129’ N, 103°15.409’ E) at 102 m in elevation. Adult male LSUHC 9376 collected on 8 September 2009 by Chan, K. O. and L. Lee Grismer on Pulau Tenggol, Terengganu, Peninsular Malaysia (04°48.111 N, 103°40.478’ E) at 83 m in elevation. Adult males (LSUHC 10710–11) collected on 24 June 2008 by Chan Kin Onn at 1030 hrs from Bukit Hangus, Pahang, Peninsular Malaysia (04°16.142’ N, 102°13.370’ E) at 10 m in elevation. Adult male LSUHC 10454 collected on 2 June 2011 by Evan S. H. Quah from the Nee Soon Swamp, Singapore (01°48.40 N, 103°49.41 E) at 20 m in elevation. Diagnosis. Maximum SVL 60.0 mm; 10 or 11 supralabials; 7–10 infralabials; keeled ventrals; no precloacal pores; moderately prominent dorsal tubercles; 17–25 paravertebral tubercles; dorsal body tubercles generally randomly arranged; tubercles absent to weak on flanks; caudal tubercles encircling tail; no tubercles in lateral caudal furrows; ventrolateral and lateral rows of caudal tubercles present; subcaudals keeled; no single, median row of enlarged, subcaudal scales; one or two postcloacal tubercles on either side of base of tail; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; 27–33 subdigital lamellae on fourth toe; regenerated tail yellow in males; posterior portion of original tail in males black (Tables 6,7).

FIGURE 61. Cnemaspis peninsularis sp. nov. Upper left: adult male (LSUDPC 962) from Pulau Tioman, Pahang, Peninsular Malaysia in the light color pattern phase sleeping on the underside of a leaf. Photograph by LLG. Upper right: adult male (LSUDPC 6703) from Gunung Ledang, Johor, Peninsular Malaysia with a regenerated tail exhibiting a caudal display. Photo by ESHQ. Lower right: juvenile (LSUDPC 964) from Pulau Tioman in the light color pattern phase sleeping on the top side of a leaf. Lower left: forest habitat of C. peninsularis sp. nov. on Pulau Tioman. Photographs by LLG.

Description of holotype. Gravid female; SVL 55.2 mm; head oblong in dorsal profile, moderate in size (HL/ SVL 0.27), somewhat narrow (HW/SVL 0.16), flattened (HD/HL 0.37), distinct from neck; snout short (ES/HL 0.47), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum weakly keeled, slightly raised, same size as similarly shaped scales on occiput; low, supraorbital ridges; very weak frontorostral sulcus; canthus rostralis not very discernable; eye large (ED/HL 0.19); extra-brillar fringe scales

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 119 155 largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 75% divided by longitudinal groove; rostral bordered posteriorly by two large supranasals and external nares, and laterally by first supralabials; 10R,L raised supralabials decreasing in size posteriorly; 8R,9L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, concave medially, bordered posteriorly by two large, rectangular, lateral postmentals of similar size and one smaller azygous scale; gular scales raised, smooth; throat scales larger, raised, weakly keeled. Body slender, elongate (AG/SVL 0.48); small, keeled, dorsal scales generally equal in size throughout body, intermixed with larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail; tubercles absent from lower flanks, moderate in size; 22 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales slightly larger than dorsals; no precloacal pores; forelimbs moderately long, slender (FL/SVL 0.19); dorsal scales of brachium raised, keeled; dorsal scales of forearm raised, keeled; ventral scales of brachium smooth, raised, juxtaposed; ventral scales of forearm weakly keeled, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout digit; interdigital webbing absent; fingers increase in length from first to fourth with fourth longer than fifth; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.22); dorsal scales of thigh keeled, raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales raised, keeled, juxtaposed, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally but wider distally throughout digit; interdigital webbing absent to weak; toes increase in length from first to fourth with fourth being slightly longer than fifth; 31 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales flat anteriorly, keeled, juxtaposed; deep middorsal and lateral caudal furrows; subcaudal scales keeled; no median row of enlarged keeled subcaudal scales; caudal tubercles encircle tail; tubercles absent from lateral furrows. Color pattern in life. Dorsal ground color yellowish; head and body overlain with irregularly shaped pattern of interconnected brownish markings highlighting white to yellowish irregularly shaped blotches; small, black, elongate medial marking on nape followed by similarly colored, paired, elongate, paravertebral markings extending to midway down body; similar vertebral markings extend from midbody to base of tail transforming into poorly defined, diffuse, dark caudal bands; limbs bearing a faint, brownish banding pattern becoming more evident distally; digits banded; ventral ground color beige; faint, darker reticulum on belly; posterior subcaudal region dark. Variation. The type series shows a modest array of color pattern variation that is not necessarily related to substrate matching although this species does tend to show differences in overall hue with respect to substrate (see Geographic variation below). LSUHC 5731 and 6253 match the holotype in general coloration and pattern although both lack the larger dark markings and appear less mottled overall. Both differ from the holotype in having a dark belly with distinct white spots as do LSUHC 10454 and 10710–11. The other specimens show a modest degree of belly mottling. LSUHC 10710–11 have a much darker color pattern that highlights the overlying lighter markings as well as a distinct, thin, whitish, nuchal loop. A distinct nuchal loop also occurs in LSUHC 4756, 8210, 8966, and 10454. LSUHC 9376 has a faded nearly unicolor dorsal pattern bearing only faint dark and light spots and LSUHC 8126 has a unicolor regenerated tail. Sexual dimorphism in caudal coloration is discussed below (see Comparisons). Meristic and mensural variation is listed in Table 14 and various color patterns can be seen in Figures 60 and 61 and in Grismer (2011a). Distribution. Cnemaspis peninsularis sp. nov. ranges as far north as Bukit Hangus, Pahang on the peninsula and to Pulau Tenggol off the east coast, and southward to Singapore (Grismer 2011a: Fig. 4). In the Seribuat Archipelago, Grismer et al. (2006) reported C. peninsularis sp. nov. from the islands of Aceh, Babi Besar, Babi Hujung, Ibol, Sembilang, Seribuat, Sibu, Sibu Tenggah, Tinggi, Tioman, and Tulai (Fig. 4). Natural History. According to Grismer (2011a) and references therein, Cnemaspis peninsularis sp. nov. is a scansorial, diurnal gecko found on logs, tree trunks, low vegetation, and rocks (Figs. 60, 61). It is a common inhabitant of both primary and secondary, lowland and hill, dipterocarp forests and to a lesser extent peatswamps and ranges up to approximately 500 m in elevation. During the day, C. peninsularis sp. nov. is active on trees and rocks beneath closed canopy forests and does not restrict its movements to dark, shaded surfaces as is commonly seen in many other species of Cnemaspis. On Pulau Tioman, Pahang, Grismer (2011a) noted that lizards may bask in sun spots on the trunks of trees. Werner and Chou (2002) indicated C. peninsularis sp. nov. is a sit and wait predator that usually perches head down on tree trunks waiting to ambush prey. When threatened, males often curl

120 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 156 their tail over their back to display the yellow underside (Fig. 61). This display is usually exaggerated by slowly moving the tail from side to side and performed just prior to fleeing to take refuge within a rock crack or beneath exfoliating bark. At night, the ground color of C. peninsularis sp. nov. becomes nearly white and somewhat transparent, highlighting the dark dorsal spots on the body and the black tail in the males (Figs. 60, 61). During this time, lizards are commonly seen sleeping on tree trunks, leaves, rocks, and clinging to the underside of leaves as high as 10 m above the ground. Grismer (2011a) reported gravid females on several islands in the Seribuat Archipelago from March through August; during June at Gunung Ledang, Johor; and at Endau-Rompin, Johor during September. In Singapore, gravid females have been observed during August and pairs of eggs stuck to rocks and cement structures have been found during November and December (Werner & Chou 2002). Hatchlings and gravid females in Singapore have been observed during December (Werner & Chou 2002). These data suggest C. peninsularis sp. nov. breeds year round. Bullock (1966) reported finding ants, beetles, earthworms, millipedes, and soil in the stomachs of lizards from Pulau Tioman indicating that foraging takes place on the ground.

TABLE 14. Meristic and mensural character states of the type series of Cnemaspis peninsularis sp. nov. w = weak; f = female; m = male; and / = data unavailable. Meristic abbreviations are listed in the Materials and Methods. For TL, the measurement for the original portion of the tail is on the left and the measurement for the regenerated portion is on the right. G. = Gunung; E. = Endau; B. = Bukit; P. = Pulau. LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC

8965 8966 8126 8210 10710 10711 10454 9376 6213 4756 5731

holotype paratype paratype paratype paratype paratype paratype paratype paratype paratype paratype G. G. E. E. B. B. P. P. P. Ledang Ledang Rompin Rompin Hangus Hangus Singapore Tenggol Tioman Tinggi P. Besar Supralabials 10 11 10 10 11 10 10 10 10 11 10 Infralabials 8 9 10 7 9 8 9 9 9 10 8 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 1 1 1 1 1 No. of precloacal pores / / / / / / / / / / / Precloacal pores continuous (1) or separated (0) / / / / / / / / / / / Precloacal pores elongate (1) or round (0) / / / / / / / / / / / No. of paravertebral tubercles 22 22 21 17 19 22 25 19 24 21 23 Tubercles linearly arranged (1) or more random (0) 0 0 / 0 w 0 0 0 w 0 w Tubercles present (1) or absent (0) on flanks w 1 / 0 w w 0 0 0 w 0 Caudal tubercles in lateral furrow (1) or not (0) 0 0 / 0 / 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 / 1 / 1 1 1 1 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 1 / 1 / 1 1 1 1 1 1 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 0 / 0 / 0 0 0 0 0 0 Subcaudals keeled (1) or not (0) 1 1 / 1 / 1 1 1 1 1 1 Single median row of keeled subcaudals (1) or not (0) 0 0 / 0 / 0 0 0 0 0 0 Caudal tubercles encircle tail (1) or not (0) 1 1 / 1 / 1 1 1 1 1 1 Enlarged median subcaudal scale row (1) or not (0) 0 0 / 0 / 0 0 0 0 0 0 Postcloacal spurs / 1 2 1 / 1 1 2 1 1 1 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 0 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 1 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 0 0 0 0 0 0 No. of 4th toe lamellae 31 29 30 28 29 29 31 29 33 27 31 Sex f m m f m m m m m m m SVL 55.2 54.7 43.4 54.6 52.6 54.6 47.3 53.0 51.3 51.1 52.1 TL 64.1 51.2 36.1 47.8 49.0 53.5 14.7 71.7 56.4 47.5 67.8 TW 5.5 5.3 3.7 5.4 4.8 5.7 4.5 5.4 5.0 5.0 5.6 FL 10.3 9.4 7.2 10.3 10.5 10.9 9.1 9.9 9.4 9.4 9.0 TBL 12.2 11.6 9.5 12.9 12.4 13.6 10.9 11.8 12.2 11.6 11.6 AG 26.3 25.2 19.1 24 22.7 25.5 20.7 22.9 22.4 22.3 21.3 HL 14.9 14 12.2 13.7 14.3 14.5 13.0 13.9 13.0 13.1 13.8 HW 8.7 8.7 7.1 8.5 8.5 8.6 7.2 8.2 8.5 7.5 8.2 HD 5.5 5.8 4.4 5.7 5.4 5.8 4.9 5.4 6.1 5.4 5.2 ED 2.9 2.9 2.4 3.0 3.0 2.9 2.7 2.9 2.9 2.9 2.6 EE 3.2 3.9 3.0 3.5 3.7 4.0 2.9 3.7 3.7 3.5 3.3 ES 7.0 6.6 5.8 6.7 6.7 7.1 5.9 6.9 6.6 6.5 6.9 EN 5.2 4.8 4.3 5.3 5.2 5.6 4.4 5.5 5.3 5.1 5.3 IO 3.2 1.3 2.6 1.3 2.8 1.3 2.6 1.4 3.0 2.9 3.0 EL 1.7 1.4 1.2 1.5 1.7 1.8 1.2 1.3 1.5 1.4 1.5 IN 1.3 1.3 1.1 1.3 1.4 1.4 1.3 1.4 1.3 1.3 1.2

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 121 157 Etymology. The specific epithet peninsularis is an adjective in reference to the distribution of this species being restricted to Peninsular Malaysia and Singapore and their adjacent islands. Geographic variation. Geographic variation in Cnemaspis peninsularis sp. nov. does not show geographically related trends as seen in some other species of lizards from Peninsular Malaysia (Grismer 2011a) but there is some noteworthy localized variation in some populations. Lizards from Pulau Tioman, Pahang generally have a more boldly marked abdomen than lizards from populations of Peninsular Malaysia. This is especially true in adult males. In extreme cases, the bellies of some males may be dark brown with white spots and that pattern may extend onto the undersides of the hind limbs. The dorsal pattern of lizards from Pulau Ibol, Johor is nearly unicolor brown. A similar pattern occurs in lizards from Sungai Lembing, Pahang except that lizards from here maintain the large, black, elongate dorsal blotches and appear superficially similar to C. baueri of Pulau Aur, Johor.

FIGURE 62. Maximum likelihood phylogram (–InL 73957.608688) of Cnemaspis peninsularis sp. nov. based on the mitochondrial gene ND2 and the geographic distribution of the specimens sampled. Closed black circles are nodes with BBP values and ML bootstrap values > 0.95 and 70, respectively.

Comparisons. Cnemaspis peninsularis sp. nov. is a member of the Southern Sunda clade which includes C. limi, C. nigridia, C. paripari, C. kendallii, C. sundainsula sp. nov., C. pemanggilensis, C. baueri, C. mumpuniae sp. nov. and C. bidongensis. Within this clade, it is part of an unresolved polytomy that composes C. kendallii, C. sundainsula sp. nov., C. pemanggilensis, C. mumpuniae sp. nov., C. bidongensis, C. peninsularis sp. nov., and C. baueri of the kendallii group (Fig. 2). Cnemaspis peninsularis sp. nov. is easily separated from C. limi by being

122 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 158 much smaller (maximum SVL 55.2 mm versus 88.2 mm); having fewer paravertebral tubercles (17–25 versus 25–35); having keeled versus smooth subcaudal scales; the presence versus the absence of a ventrolateral row of caudal tubercles; having caudal tubercles that encircle the tail versus not having tubercles that encircle the tail; and lacking versus having white caudal tubercles. From C. paripari, C. peninsularis sp. nov. lacks precloacal pores as opposed to having them; has fewer paravertebral tubercles (17–25 versus 26–31); has as opposed to lacks tubercles on the flanks; has versus lacks a ventrolateral row of caudal tubercles; has caudal tubercles that encircle the tail versus not having tubercles encircling the tail; lacks versus has an enlarged, median subcaudal scale row; and males lack as opposed to having a yellow head, limbs, and back and the posterior one-half of the original tail being white. Within the kendallii group, C. peninsularis sp. nov. is easily distinguished from C. sundainsula sp. nov., C. pemanggilensis, and C. baueri by being much smaller (maximum SVL 55.2 mm versus 67.4–84.5 mm) and from C. sundainsula sp. nov. it is further separated by having keeled versus smooth ventral scales, and caudal tubercles that encircle the tail rather than not having such tubercles. Cnemaspis peninsularis sp. nov. is further separated from C. pemanggilensis by having fewer paravertebral tubercles (17–25 versus 30–37) and lacking as opposed to having an enlarged, median row of keeled subcaudal scales. From C. baueri, C. peninsularis sp. nov is further differentiated by lacking an enlarged, median row of keeled subcaudal scales and not having a uniform brown dorsal color pattern bearing large, elongate black blotches on the nape and in the shoulder region. Cnemaspis peninsularis sp. nov. can not be differentiated from C. kendallii (with which it was previously considered conspecific) on the basis of scale counts. However, these species do differ notably in that the abdomen of C. peninsularis sp. nov. is mottled with a diffuse, dark reticulum enclosing lighter spots whereas in C. kendallii the abdomen is beige and generally immaculate; the posterior two-thirds of the original tail in adult male C. peninsularis sp. nov. is black dorsally and ventrally and in adult male C. kendallii the tail is banded dorsally throughout its length and the subcaudal region is essentially immaculate white. The regenerated tail in adult male C. peninsularis sp. nov. is yellow and immaculate dorsally and ventrally whereas that of C. kendallii is straw colored with small black flecks dorsally and the subcaudal region is white and immaculate. Additionally, C. kendallii has a row of nearly contiguous tubercles on the lateral margings of the occipital region bordering the nape which are nearly always absent in C. peninsularis sp. nov. Another row of tubercles generally absent in C. peninsularis sp. nov. occurs immediately anterior to the shoulder region in C. kendallii. All these tubercles are usually accentuated by being white. Within the kendalli group, C. peninsularis sp. nov. is most closely related to C. bidongensis and C. mumpuniae sp. nov. (Fig. 2). It differs from C. mumpuniae in lacking a brick-red ground color and a thin, white, nuchal loop. It is differentiated further from C. bidongensis sp. nov. by lacking as opposed to having an enlarged, median, subcaudal scale row. Relationships. Cnemaspis peninsularis sp. nov. forms a polytomy with C. bidongensis from Pulau Bidong, Peninsular Malaysia and C. mumpuniae sp. nov. from Pulau Natuna Besar, Indonesia (Fig. 2). Remarks. Cnemaspis peninsularis sp. nov. is the most widely distributed species withn a genus that is composed primarily of microhabitat specialist with highly circumscribed distributions (Figs. 3, 4). One of the reasons for its relatively wide distribution is that it is a habitat generalist active both day and night on all rocky and vegetative substrates. To encompass the phylogeographic structure within C. peninsularis sp. nov., we sampled 32 specimens from throughout the extent of its distribution from Singapore in the south to Bukit Hangus, Pahang, and Pulau Tenggol, Terengganu in the north (Fig. 62) as well as from seven of the 11 islands on which it is known to occur in the Seribuat Archipelago (Fig. 62). An ND2 phylogeny shows that C. peninsularis sp. nov. is composed of three well-supported, major lineages (Fig. 62). The basal lineage is represented by a northern population from Sungai Lembing, Pahang and its sister lineage is composed of a southern population from Pulau Seribuat, Johor of the Seribuat Archipelago and its sister lineage containing the remaining populations that generally encompasses the entire range of C. peninsularis sp. nov. (Fig. 62). The overall phylogeographic structure within the latter widespread population is polytomous with little concordance between geographic distribution and phylogenetic substructuring (Fig. 62). We consider this lack of geographic clustering to be evidence of gene flow within this widely distributed, habitat generalist (see Grismer et al. [2012] and Johnson et al. [2012] for other gekkonid examples with similar phylogeographic structure from Peninsular Malaysia). Additionally, the phylogeny indicates that the presence of C. peninsularis sp. nov. in the Seribuat Archipelago may be due to sequential vicariant events associated with episodic changes is sea levels or independent dispersal events. The first event established the Pulau Seribuat population followed by the establishment of a population on Pulau Tulai. The widespread clade of insular populations in the Seribuat Archipelago may have resulted from the most recent vicariant event and being that the

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 123 159 peninsular, Gunung Berlumut population is deeply embedded within this island clade, suggests an upstream recolonization of Peninsular Malaysia may have occurred (Fig. 62). These hypotheses and others are currently being tested with fine-scaled geographic sampling and re-analysis (Wood & Grismer in prep.). The uncorrected pairwise genetic distances among the three major lineages of Cnemaspis peninsularis sp. nov. ranges from 1.1%–9.9% whereas that within the most widespread lineage (i.e. the lineage that does not include Sungai Lembing and Pulau Seribuat; Fig. 62) ranges from 1.1%–6.7% (Table 5). The distances between the Sungai Lembing and Pulau Seribuat populations to each other and all other C. peninsularis sp. nov. are commensurate with that between other species of gekkonids (see Grismer et al. 2013b) and their specific status will also be evaluated (Wood & Grismer in prep.). Additional material examined. Peninsular Malaysia: Johor: Bunker Trail ZRC 2.5602; Endau-Rompin LSUHC 7691, 8122, 8126, 8191, 8210; Gunung Ledang ZRC 2.5437–38, LSUHC 8965–67; Pulau Babi Besar LSUHC 5731–34; Pulau Babi Hujung LSUHC 5749–52; Pulau Ibol LSUHC 6380–83; Pulau Sembilang LSUHC 5244; Pulau Seribuat LSUHC 5184–87, 5198, 5211; Pulau Tinggi LSUHC 4707, 4756–57, 4765–67; Pulau Tulai LSUHC 3894, 5056–58. Pahang: Bukit Ringgit DWNP 2231; Gemas ZRC 2.1105–06; Jerantut ZRC 2.1101; Kuala Gandah DWNP 475–76; Lakum Forest Reserve DWNP 2278–79; Pulau Tioman DWNP 1833, LSUHC 3773–75, 3797, 3811, 3820, 3841, 3878–88, 4566, 4570, 4615, 4658–59, 4666; 5436, 5445–46, 5454, 5462, 5477, 5482, 6213–18, 6224, 8036; Sungai Lembing LSUHC 4954, 4958. Negeri Sembilan: Gallah Forest Reserve DWNP 2281. Selangor: Sungai Lalang DWNP 169; Ulu Gombak DWNP 1828. Singapore: ZRC 2.107–08, 2.3014, 2.3520, 2.3544, 2.4992, 2.5644, 2.5891.

Cnemaspis bidongensis Grismer, Wood, Amirrudin, Sumarli, Vazquez, Ismail, Nance, Muhammad, Mohamad, Syed, Kuss, Murdoch & Cobos, 2014 Pulau Bidong Rock Gecko Fig. 63

Holotype. Adult female (LSUHC 11455) collected on 26 August 2013 by Jacob A. Chan at 2200 hrs at 34 m from Pulau Bidong, Terengganu, Peninsular Malaysia (5°37.201 N 103°03.244 E) at 49 m in elevation. Paratype. Adult males (LSUHC 11447, 11452–54) and adult female LSUHC 11451 bear the same data as the holotype except they were collected between 1800 and 2400 hrs. Diagnosis. Maximum SVL 58.1 mm; nine or 10 supralabials; 7–9 infralabials; keeled ventral scales; no precloacal pores; 21–26 paravertebral tubercles; no tubercles on flanks; caudal tubercles encircling tail; no tubercles in lateral caudal furrows; ventrolateral caudal tubercles present anteriorly; subcaudals keeled; a median row of enlarged, keeled subcaudals; one or two postcloacal tubercles on each side; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; 26–30 subdigital lamellae on fourth toe; large, round, black blotches on nape and anterior portion of body; distinct black and white caudal bands in females (Tables 6,7). Color pattern in life (Fig. 63). Females: dorsal ground color of head, body, limbs and tail brown; a series of diffuse, yellowish lines on rostrum extend posteriorly onto frontal region; dorsal pattern on occiput consisting of a series of dull white spots surrounding a dark brown, tear-drop shaped, vertebral marking; three diffuse, dark brown, postorbital stripes radiate from eyes; dorsal pattern of neck and body consists of a vertebral series of six dull-white blotches extending to base of tail paralleled by similar blotches on flanks; a series of seven diffuse dark brown blotches extend from side of neck along flanks to base of tail on each side of body; intervening area between all body blotches consists of a network of dark and light mottling that extends onto the limbs; nine white caudal bands infused with faint black speckling encircle tail; interband areas bear black mottling; ventral surfaces of head body and limbs beige to dull yellow with weak black stippling in each scale; anterior gular region yellow; subcaudal region bearing white, irregularly shaped bands; small, dark, elongate blotch on mental and medial postmental. Males: overall yellowish to dull-orange dorsal pattern on the head, body and tail and lack the dull-white blotching seen in females yet retain the darker blotching pattern. The caudal pattern is banded but the light bands are not white as in the females and the dark bands are dark brown as opposed to black. The ventral pattern of males is similar to that of females except that the lateral margins of the abdomen tend to near dark network enclosing small, lighter spots. Also, subcaudal banding is faint. Distribution. Cnemaspis bidongensis is presumed to be endemic to Pulau Bidong, Terengganu, Peninsular

124 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 160 Malaysia (Grismer et al. 2014; Fig. 4) being that it has not been found in any other archipelago or any other island in the Bidong Archipleago (Vazquez et al. 2014.). Natural History. Grismer et al. (2014) noted that Cnemaspis bidongensis occurs in secondary, coastal forest and is widespread throughout the island. During the Vietnamese refugee period from May 1975 to October 1991, the island’s primary forest was severely degraded by cutting. During this time, as many as 250,000 people fleeing the communist take over of southern Vietnam spent time on Pulau Bidong and in June 1979 it was considered the most heavily populated place on earth (http://en.wikipedia.org/wiki/Bidong_Island). Although this had a catastrophic effect on the native forest, C. bidongensis was able to survive because it is not a microhabitat specialist as are many other species of Cnemaspis (Grismer & Ngo 2007; Grismer & Chan 2009; Grismer et al. 2010a,b; 2013a; Grismer 2011a; Wood et al. 2013). During the course of our fieldwork, lizards were observed day and night on both granite rocks and vegetation (Fig. 63). Lizards were wary, swift, agile and would seek shelter at the slightest provocation. During the day, lizards would often jump from rocks to nearby trees and escape by ascending 3–5 meters up the trunk—a behavior not observed in any other species of Cnemaspis. Lizards would also avoid capture by retreating into rock cracks. During the evening, lizards were commonly seen on rocks, branches, and leaves where they appeared to be sleeping. When aroused, many would drop to the forest floor from as high as 1.5 meters and escape into the leaf litter—a behavior also uncharacteristic of Cnemaspis. Hatchlings and juveniles were not observed and the presence of gravid females carrying two eggs suggests that July is the beginning of the reproductive season. Relationships. Cnemaspis bidongensis froms a polytomy with C. peninsularis sp. nov. from Peninsular Malaysia and C. mumpuniae sp. nov. from Pulau Natuna Besar, Indonesia (Fig. 2).

FIGURE 63. Cnemaspis bidongensis from Pulau Bidong, Terengganu, Peninsular Malaysia. Upper left: adult female (LSUDPC 8090) in the dark color pattern phase. Upper right: adult male (LSUDPC 8096) in the dark color pattern phase. Lower left: ventral view of adult male (upper) and (lower) female (LSUDPC 8108). Lower right: granite boulder and forest microhabitat on Pulau Bidong. Photographs by LLG.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 125 161 Species Incertae Sedis

We were unable to acquire tissue for five previously described and one newly described species herein: Cnemaspis laoensis Grismer; C. punctatonuchalis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya; C. vandeventeri Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya; C. kamolnorranathi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya; C. dringi Das & Bauer, and C. sundagekko sp. nov. Thus, their phylogenetic relationships remain unknown. However, based on morphology and the fact that the smaller clades (i.e., Ca Mau and Pattani) and species groups are restricted to well-circumscribed regions with virtually no overlap in distribution (except for the Pattani and Northern Sunda clades) we can use these data to hypothesize to which clade and/or group each species may belong.

Cnemaspis laoensis Grismer, 2010 Lao Rock Gecko Fig. 64

Holotype. THNHM 12433. Type locality: “Dong Phu Vieng National Protected Area, Savannakhet, Laos”. Diagnosis. Maximum SVL 40.9 mm; nine supralabials; seven infralabials; smooth ventral scales; no precloacal pores; 22 paravertebral tubercles; body tubercles randomly arranged, present on flanks; tubercles present in lateral caudal furrows; no ventrolateral or lateral caudal row of tubercles; caudal tubercles do not encircle tail; subcaudals smooth, bearing a slightly enlarged median row; two or three postcloacal tubercles; no enlarged femoral, subtibial or submetatarsal scales; subtibials keeled; and 29 subdigital fourth toe lamellae (Table 6). Cnemaspis laoensis lacks the diagnostic color characteristics of other Southern Indochina clade species.

FIGURE 64. Adult female holotype of Cnemaspis laoensis (THNHM 12433) from Dong Phu Vieng National Protected Area, Savannakhet, Laos. Photograph by LLG.

126 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 162 Color pattern (Fig 64). Dorsal ground color of head, body, limbs and tail pale brown; rostrum and top of head bearing diffuse, faint, brown markings; postorbital stripes absent; light-colored, subelliptical, paired, paravertebral markings extending from nape to base of tail, continuing onto tail as wide, diffuse, light-colored, caudal bands separated by thinner, dark-brown bands; dark-brown markings within interspaces between light markings; large, light-colored markings on flanks; forelimbs and hind limbs mottled; all ventral surfaces beige bearing small, black stipples in each scale. Distribution. Cnemaspis laoensis is known only from the type locality at Dong Phu Vieng, Savannakhet, along the Laotian portion of the Ho Chi Minh Trail (Fig. 1). Natural History. Nothing is known of the natural history of Cnemaspis laoensis. However, Grismer (2010) hypothesized that on the basis of having smooth ventral and subcaudal caudal scales, this species is a likely karst- dweller. All other karst dwelling, microhabitat specialists share this combination of characters although its prominent dorsal tuberculation would suggest otherwise. Additionally, there are numerous outcroppings of karst tower formations in the general area of the type locality. Relationships. The distribution or Cnemaspis laoensis in southern Laos (Fig. 1) would suggest that it could potentially belong in the Ca Mau clade, the caudanivea group of the Southern Indochina clade of southern Vietnam or the chanthaburiensis group of southern Thailand and southwestern Cambodia of the Southern Indochina clade. The fact that this species has a small SVL (40.9 mm); dense, prominent, dorsal tuberculation; lacks enlarged femoral, subtibial and submetatarsal scales; has a dorsolateral row of caudal tubercles; and tubercles in the lateral caudal furrow would likely preclude it from being a member of the Ca Mau clade (Table 6). Its dense arrangement of prominent dorsal tubercles; lack of a ventrolateral row of caudal tubercles; and lack of a dark, mid-gular stripe would likely preclude it from being a member of the caudanivea group (Table 6). It differs from members of the chanthaburiensis group only in lacking a lateral row of caudal tubercles and thus its inclusion in this group would be the most likely (Table 6). However, given that C.laoensis is separated from the nearest member of the chanthaburiensis group in Khao Ang Ru Ni, Chachoengsao Province in southeastern Thailand by approximately 600 km, two mountain ranges, and the Tonle Sap Basin would indicate that it will likely prove to be in its own species group within the Southern Indochina clade. For now, we tentatively place it in the chanthaburiensis group. Material examined. Laos: Savannakhet, Dong Phu Vieng National Protected Area THNHM 12433 (holotype).

Cnemaspis punctatonuchalis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Spotted-neck Rock Gecko Fig. 65

Holotype. THNHM 2001. Type locality: “Thap Sakae District, Prachuap Khirikhan Province, Thailand” at approximately 70 m in elevation. Diagnosis. Maximum SVL 49.6 mm; eight supralabials; seven or eight infralabials; smooth ventral scales; no precloacal pores; 24–27 paravertebral tubercles; body tubercles semi-linearly arranged, present on flanks; tubercles absent from lateral caudal furrows; ventrolateral caudal row of tubercles present anteriorly; lateral caudal row of tubercles present; caudal tubercles do not encircle tail; subcaudals smooth, bearing an enlarged median scale row; 1–3 postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials smooth; 29–31 subdigital fourth toe lamellae; white ocelli on brachia and side of neck in males; throat and subcaudal region orange in males (Tables 6,7). Color pattern in life (Fig. 65). Dorsal ground color of head, body, limbs and tail light brown; top of head bearing large, diffuse, light and dark-colored markings giving it a mottled appearance; postorbital stripes absent; large, whitish markings on side of head forming a reticulum that extends ventrally onto throat; three, radiating, elongate blotches on occiput bordering the anterior margin of a large, white spot; lower sides of neck black, enclosing a large, white ocellus; black neck patch edged posteriorly by white antebrachial and brachial markings; lightly colored, paravertebral, butterfly-shaped markings between forelimb insertions and base of tail; markings continue onto tail to form light-colored bands; patches of enlarged tubercles on flanks white; other tubercles on body dark or light-colored; limbs generally uniform brown bearing elbow and knee patches; all ventral surfaces uniform beige with fine, dark stippling in some scales. Sexual dimorphism is distinct. Adult males have a

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 127 163 brownish-green head with a yellow neck that accentuates the black, neck patch bearing the whitish ocelli. The body and limbs are grayish green and the body bears a series of alternating dark and light, paravertebral blotches. The ground color of the tail is deep-yellow and overlain with lighter, yellow bands. Adult females have a greyish dorsal ground color overall that is overlain by a prominent series of light and dark blotches on the head and a large, light- colored, central nape blotch. Alternating dark and light, paravertebral blotches occur on the body and transform into a lighter banding pattern on the tail. The limbs are somewhat banded distally and spotted proximally. The tubercular patches on the flanks are smaller in females than in males.

FIGURE 65. Cnemaspis punctatonuchalis from Thap Sakae District, Prachuap Khirikhan Province, Thailand. Upper left: adult male (LSUDPC 8087) in the dark color pattern phase. Upper right: adult female (LSUDPC 8088) in the dark color pattern phase. Lower: granite boulder microhabitat at Thap Sakae. Photographs by MS.

Distribution. Cnemaspis punctatonuchalis is known only from the type locality of the district of Thap Sakae, Prachuap Khiri Khan Province, Thailand (Grismer et al. 2010a; Fig. 3). Natural history. Specimens have only been observed at night on granite boulders in lowland forest (Grismer et al. 2010a; Fig. 65). Relationships. The distribution of Cnemaspis punctatonuchalis in Peninsular Thailand would align it with members of the siamensis group (Fig. 3). Like the northern sister species of this group C. huaseesom and C. siamensis, C. punctatonuchalis occurs north of the Isthmus of Kra and lacks the light colored, prescapular crescent that diagnoses the monophyletic group composed of C. chanardi, C. omari sp. nov., and C. roticanai that occurs south of the Isthmus of Kra, suggesting it may be more closely related to the northern species (Table 6). Material examined. Thailand: Prachuap Khirikhan Province, Thap Sakae District THNHM 1899, 2001; Thap Sakae District, Hauy Yang ZMKU Rep-000314 (type series).

128 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 164 Cnemaspis vandeventeri Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Vandeventer’s Rock Gecko Fig. 66

Gonatodes siamensis Smith, 1925:22? Cnemaspis siamensis Smith, 1935:72; Taylor, 1963:743 Cnemaspis siamensis (?) Pauwels et al., 2000:129

Holotype. THNHM 8261. Type locality “Khlong Naka Wildlife Sanctuary (9°26.0N, 98° 35.0E), Kapur District, Ranong Province; Thailand” at approximately 11 m in elevation. Diagnosis. Maximum SVL 44.7 mm; eight or nine supralabials; 7–9 infralabials; keeled ventral scales; four pore-bearing precloacal scales with round pores; 25–29 paravertebral tubercles; body tubercles randomly arranged, absent from flanks; tubercles absent from lateral caudal furrows; no ventrolateral row of caudal tubercles; lateral row of caudal tubercles present; caudal tubercles do not encircle tail; subcaudals keeled, bearing a weakly keeled, enlarged median scale row; 1–3 postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials keeled; 24–28 subdigital fourth toe lamellae; yellowish, prescapular crescent; gular region, throat, pectoral region, underside of limbs, belly, and subcaudal region orange (Tables 6,7).

FIGURE 66. Cnemaspis vandeventeri from Phuket Island, Phuket Province, Thailand. Upper right: adult male (LSUDPC 4362) in the dark color pattern phase. Photograph by H. Bringsoe. Lower right: juvenile (LSUDPC 8541) from Ranong Province in the light color pattern phase. Left: vegetative microhabitat on Phuket Island with a potential predator Dryocalamus subannulatus Duméril, Bibron & Duméril, 1854 (striped pattern phase). Photographs by MS.

Color pattern (Fig. 66). Dorsal ground color of head, body, limbs and tail brown; top of head bearing small, faint, brown markings and postorbital stripes; series of small, light-colored, vertebral blotches extend from nape to level of hind limb insertions; yellowish, prescapular crescent followed by a series of irregularly shaped, light- colored blotches on flanks; limbs faintly mottled with diffuse, dark markings; all ventral surfaces cream-colored, immaculate except for small, individual stipples in each scale. Distribution. Cnemaspis vandeventeri is restricted to the west side of the Tenasserim Mountains and the contiguous Phuket Mountains along the west coast of southern Peninsular Thailand (Grismer et al. 2010a; Fig. 3). It ranges from the Khlong Naka Wildlife Sanctuary in the north, southward approximately 58 km to Khlong Had Sompen, Ranong and onto Phuket Island. Pauwels et al. (2000) collected two specimens (MNHN 1999.7707–08) from Phang-Nga Wildlife Breeding Station, Phang-Nga located west of the Phuket Mountains that they referred to

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 129 165 as C. siamensis which could have also been C. vandeventeri. Unfortunately, the specimens could not be located (P. David, in lit. 2009). Cnemaspis vandeventeri may extend farther north along the western flanks of the Tenasserim Mountains into Myanmar. Natural history. Cnemaspis vandeventeri has been observed on or within vegetation at night in lowland vegetation as well as on granite rocks suggesting it is a habitat generalist (Fig. 66). Relationships. The distribution of Cnemaspis vandeventeri in Peninsular Thailand would align it with members of the siamensis group (Fig. 3). Unlike the northern sister species of this group C. huaseesom and C. siamensis, C. vandeventeri is restricted to the northwestern edge of the Isthmus of Kra and has the yelowish, prescapular crescent that diagnoses the monophyletic lineage composed of C. chanardi, C. omari sp. nov., and C. roticanai that occurs south of the Isthmus Kra, suggesting it may be more closely related to these species (Table 6). Material examined. Thailand: Ranong Province, Kapur District, Khlong Naka Wildlife Sanctuary THNHM 8260–1; Muang District, Ranong Province, Khlong Had Sompen, CUMZ-R- 2009,6,24–11. These represent the type series.

FIGURE 67. Cnemaspis kamolnorranathi from the Petchphanomwat Waterfall, in Tai Rom Yen National Park, Surat Thani Province, Thailand. Upper right; adult male (LSUDPC 8530) in the light color pattern phase. Lower right: juvenile (LSUDPC 8533) in the dark color pattern phase. Right: habitat at Petchphanomwat Waterfall. Photographs by MS.

Cnemaspis kamolnorranathi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010 Kamolnorranath’s Rock Gecko Fig. 67

Cnemaspis siamensis Grismer, Chan, Nurolhuda, & Sumontha, 2008a:54

Holotype. THNHM 15908. Type locality “Petchphanomwat Waterfall, in Tai Rom Yen National Park, Ban Nasan District, Surat Thani Province (8°56.88’N 99°31.82’E)”, Thailand at 5 m in elevation. Diagnosis. Maximum SVL 37.8 mm; eight or nine supralabials; seven or eight infralabials; smooth to weakly keeled ventral scales; six or seven contiguous, pore-bearing, precloacal scales with elongate pores; 19–24

130 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 166 paravertebral tubercles; body tubercles semi-linearly arranged, present on flanks; tubercles present in lateral caudal furrows; ventrolateral row of caudal tubercles absent; lateral row of caudal tubercles; caudal tubercles do not encircle tail; subcaudals keeled, bearing a median row of weakly enlarged scales; one or two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials may or may not be keeled; 24–28 subdigital fourth toe lamellae; light-colored prescapular crescent variably present (Tables 6,7). Color pattern in life (Fig. 67). Dorsal ground color of head, body, limbs and tail brown to pale-yellow; rostrum bearing dark and light irregular markings; occiput darker than top of head; postorbital stripping faint to prominent; light vertebral blotch on nape followed by 4–6, lightly colored, butterfly-shaped, vertebral blotches on body which may fade immediately anterior to level of groin; white markings weakly edged in darker coloration; no light-colored bars on flanks; light-colored prescapular crescent variably present; limbs bearing dark and light- colored mottling faintly resembling a banding pattern; ventral surfaces uniformly beige with faint, black stippling in all scales; distinct dark and light color phases. Distribution. Cnemaspis kamolnorranathi is known only from the type locality at Petchphanomwat Waterfall, in Tai Rom Yen National Park, Kanchanadid District, Surat Thani Province and Tham Khao Sonk hill, Thachana District, Surat Thani Province, Thailand (9°34’N 99°10’E), approximately 110 km to the north (Grismer et al. 2010a; Fig. 3). Natural history. Grismer et al. (2010a) noted that Cnemaspis kamolnorranathi expresses a wide range of substrate utilization. Lizards have been found on karst, beneath rocks, and on vegetation and buildings during evening hours in lowland forests (Fig. 67). Remarks. Grismer et al. (2010a) noted that the relatively wide separation (~110 km) between the Petchphanomwat Waterfall and Tham Khao Sonk suggests there are probably undiscovered, geographically intervening populations of Cnemaspis kamolnorranathi in the appropriate habitat separating these two localities. Unlike other species of Cnemaspis, C. kamolnorranathi shows intrapopulational variation in the degree of keeling of the ventral and subtibial scales suggesting C. kamolnorranathi may be composed of more than one species. Relationships. The distribution of Cnemaspis kamolnorranathi in Peninsular Thailand would align it with members of the siamensis group (Fig. 3). Unlike the northern sister species of this group C. huaseesom and C. siamensis, C. kamolnorranathi is restricted to the Isthmus of Kra but has a variably present light-colored, prescapular crescent that diagnoses a monophyletic group composed of C. chanardi, C. omari sp. nov., and C. roticanai that occurs south of the Isthmus Kra (Table 6). This would suggest C. kamolnorranathi may be more closely related to southern species. Material examined. Thailand: Surat Thani Province, Ban Nasan District, Tai Rom Yen National Park, Petchphanomwat Waterfall THNHM 15908, PSUZC-RT 2010.52, KZM 006; Thachana District, Tham Khao Sonk hill CUMZ-R 2009,6,24-3. These specimens represent the type series.

Cnemaspis dringi Das & Bauer, 1998 Dring’s Rock Gecko Fig. 68

Holotype. FMNH 148588. Type locality “Labang Camp (03° 20'N; 113°29'E), Bintulu District, Fourth Division, Sarawak, East Malaysia, Borneo” at approximately 30 m in elevation. Diagnosis. Maximum SVL 46.4 mm; 11 supralabials; nine infralabials; keeled ventral scales; five or six discontinuous, pore-bearing, precloacal scales with round pores; 25–27 paravertebral tubercles; body tubercles randomly arranged, absent from flanks; tubercles absent from lateral caudal furrows; no ventrolateral row of caudal tubercles; lateral row of caudal tubercles present; caudal tubercles do not encircle tail; subcaudals keeled; enlarged median row of subcaudals present; two postcloacal tubercles on each side of tail base; no enlarged femoral, subtibial or submetatarsal scales; subtibials keeled; 32–35 subdigital fourth toe lamellae; distinct, whitish spots on a dark flank (Tables 6,7). Color pattern (Fig. 68). Dorsal ground color pale brown overlain with a vertebral series of irregularly shaped, light-colored markings flanked by a linear, dark, paravertebral series of markings extending from nape to beyond base of tail; two dark-brown, postorbital stripes extending to ear opening; upper surface of limbs bearing dark bands; ventral surfaces pale-brown with scattered clusters of off-white scales, manifesting a very weakly mottled appearance; flanks dark-brown bearing distinct, whitish spots of varying size.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 131 167 FIGURE 68. Type series of Cnemaspis dringi from Sarawak, East Malaysia. Upper: adult male (FMNH 148588; upper) from Fourth Division, Bintulu District, Labang Camp and adult male (FMNH 221478: lower) from seventh Division, Belaga District. Lower: FMNH 221448 showing diagnostic spotting on flank.

Distribution. Cnemaspis dringi is known only from Labang Camp, Bintulu District, Fourth Division and Sungai Segaham, Belaga District, Seventh Division, Sarawak, East Malaysia (Das & Bauer 1998; Fig. 4). Natural history. According to Das and Bauer (1998), the only known aspect of this species’ natural history is that the paratype from Sungai Segaham was taken from a log. Relationships. The distribution of Cnemaspis dringi in East Malaysia would suggest it belongs within the Southern Sunda clade (Fig. 4). Having precloacal pores like all species of the nigridia group of East Malaysia differentiates C. dringi from all species in the kendallii group as well as C. limi. Like all species of the nigridia group, C. dringi lacks tubercles encircling the tail, which further separates it from the kendallii group. These data (Tables 6,7) would suggest that C. dringi is most closely related to the East Malaysian nigridia group. Material examined. Malaysia: Sarawak, Fourth Division, Bintulu District, Labang Camp FMNH 148588; seventh Division, Belaga District FMNH 221478. These specimens represent the type series.

132 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 168 FIGURE 69. Type series of Cnemaspis sundagekko sp. nov. from Pulau Siantan, Anambas Archipelago, Riau Province, Indonesia. Upper: adult male holotype USNM 26549 (far left) and adult male paratypes USNM 26547–48 middle and right, respectively. Lower: paratypes ZRC. 2.1109–11 from left to right, respectively.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 133 169 Cnemaspis sundagekko sp. nov. Anambas Rock gecko Fig. 69

Gonatodes kendalli Smedley 1928:76 Cnemaspis cf. kendallii Leong, Grismer & Mumpuni 2003:168

Holotype. Adult male USNM 26549 collected during September of 1899 by W. Abbott on Pulau Siantan, Anambas Archipelago, Riau Province, Indonesia (03°09.01 N, 106°14.03 E). Elevation unkown but less than 400 m, the maximum height of the island. Paratypes. All paratypes are from Pulau Siantan. Adult males USNM 26547–48 bear the same data as the holotype. Adult males ZRC 2.1109–10 and adult female ZRC 2.1111 were collected on 5 September 1925 by F. N. Chasen. Diagnosis. Cnemaspis sundagekko sp. nov. differs from all other Southeast Asia species of Cnemaspis in having the unique combination of adult males reaching 65.6 mm SVL, adult females reaching 68.0 mm SVL; 11–13 supralabials; 8–11 infralabials; keeled ventrals; no precloacal pores; moderate to prominent dorsal tubercles; 20–25 paravertebral tubercles; dorsal body tubercles generally randomly arranged; tubercles absent to weak on flanks; caudal tubercles encircle tail; no tubercles in lateral caudal furrows; ventrolateral and lateral rows of caudal tubercles present; subcaudals keeled; a sinlge, median row of enlarged, keeled subcaudals posteriorly; two or three postcloacal tubercles on either side of base of tail; no enlarged femoral, subtibial, or submetatarsal scales; subtibials keeled; 31–38 subdigital lamellae on fourth toe; large, dark, round spots on nape and anterior portion of body; dorsal caudal tubercles white. These characters are scored across all species in Tables 6 and 7. Description of holotype. Male; SVL 55.2 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.27), somewhat narrow (HW/SVL 0.17), flattened (HD/HL 0.37), distinct from neck; snout short (ES/HL 0.50), slightly concave in lateral profile; postnasal region constricted medially, flat; scales of rostrum smooth, raised, larger than scales on occiput; low, supraorbital ridges; no frontorostral sulcus; canthus rostralis not very discernable; eye large (ED/HL 0.19); extra-brillar fringe scales largest anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, nearly completely divided by longitudinal groove; rostral bordered posteriorly by two large supranasals and external nares, and laterally by first supralabials; 13R,11L raised supralabials decreasing in size posteriorly; 10R,9L infralabials, decreasing in size slightly posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered posteriorly by small, granular, postnasal scales; mental large, triangular, flat, bordered posteriorly by two large, rectangular, lateral postmentals of similar size and one smaller azygous scale; gular scales raised, weakly keeled; throat scales larger, raised, keeled. Body moderate in stature (AG/SVL 0.41); small, keeled, dorsal scales generally equal in size throughout body, intermixed with larger, multicarinate tubercles more or less randomly arranged; tubercles extend from occiput to base of tail, moderate in size; tubercles absent from lower flanks; 25 paravertebral tubercles; pectoral and abdominal scales raised, keeled, not elongate, same size throughout; abdominal scales same size as dorsals; no precloacal pores; forelimbs moderately long, slender (FL/SVL 0.20); dorsal scales of brachium raised, keeled; dorsal scales of forearm raised, keeled; ventral scales of brachium keeled, raised, juxtaposed; ventral scales of forearm weakly keeled, raised, juxtaposed; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout digit; interdigital webbing absent; fingers increase in length from first to fourth with fourth longer than fifth; hind limbs slightly longer and thicker than forelimbs (TBL/SVL 0.22); dorsal scales of thigh keeled, slightly raised, juxtaposed; scales of anterior margin of thigh keeled; ventral scales of thigh keeled; subtibial scales raised, keeled, juxtaposed, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout digit; interdigital webbing absent to weak; toes increase in length from first to fourth with fourth being slightly longer than fifth; 31 subdigital lamellae on fourth toe; caudal scales arranged in segmented whorls; dorsal caudal scales flat anteriorly, keeled, juxtaposed; deep middorsal and lateral caudal furrows; subcaudal scales keeled; median row of enlarged, keeled, subcaudal scales posteriorly; caudal tubercles encircle tail; tubercles absent from lateral furrows; two enlarged, postcloacal tubercles on lateral surface of hemipenal swellings at base of tail. Color pattern (Fig. 69). The type material was collected between 1899 and 1925 and is currently devoid of pattern save for some very faint markings. Smedley (1928) provides a brief color description taken from notes

134 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 170 made in the field by Mr. F. N. Chasen in 1925 as follows: “Greyish-green above, blotched with brown; brighter green before eyes. Light rings on tail greyish-white; dark rings greenish brown. Below whitish. Distinct large brown spots on head and shoulders. Chiefly dark brown above; large oval spots on neck, nape and shoulders and no green anywhere.” From these descriptions and what can be discerned from the type material, Cnemaspis sundagekko sp. nov. has a series of medium-sized dark dorsal blotches alternating with similarly sized lighter blotches. The large, keeled, spinose caudal tubercles are accentuated by being white. Variation. No useful color pattern variation can be gleamed from the faded condition of the type material. Meristic and mensural variation is listed in Table 15.

TABLE 15. Meristic and mensural character states of the type series of Cnemaspis sundagekko sp. nov. w = weak; f = female; m = male; post = posterior; and / = data unavaiable. Meristic abbreviations are listed in the Materials and Methods.

ZRC ZRC ZRC USNM USNM USNM 2.1109 2.111 2.1111 26547 26548 26549

paratype paratype paratype paratype paratype holotype

SVL 52.3 65.6 68 64.4 61.2 65.2 Supralabials 11 12 11 12 11 13 Infralabials 9 8 9 8 11 10 Ventral scales keeled (1) or not (0) 1 1 1 1 1 1 No. of precloacal pores 0 0 / / 0 / Precloacal pores continuous (1) or separated (0) / / / / / / Precloacal pores elongate (1) or round (0) ////// No. of paravertebral tubercles 22 25 25 23 20 25 Tubercles linearly arranged (1) or more random (0) w w 0 w w w Tubercles present (1) or absent (0) on flanks w w 0 0 0 0 Caudal tubercles in lateral furrow (1) or not (0) 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 1 1 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 1 1 Caudal tubercles restricted to a single paravertebral row on 0 0 0 0 0 0 each side (1) or not (0) Subcaudals keeled (1) or not (0) 1 1 1 1 1 1 Single median row of keeled subcaudals (1) or not (0) post post post post post post Caudal tubercles encircle tail (1) or not (0) 1 1 1 1 1 1 Enlarged median subcaudal scale row (1) or not (0) post post post post post post Postcloacal spurs 2 3 / 3 2 2 Enlarged femoral scales present (1) or absent (0) 0 0 0 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 Subtibial scales keeled (1) or not (0) 1 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 0 0 0 No. of 4th toe lamellae / 33 38 33 31 31 Sex m m f m m m SVL 52.3 65.6 68.0 64.4 61.2 65.2 TL /// 77.0 / 81.5 TW 4.8 6.7 6.7 7.3 5.8 6.3 ...... continued on the next page

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 135 171 TABLE 15. (Continued) ZRC ZRC ZRC USNM USNM USNM 2.1109 2.111 2.1111 26547 26548 26549

paratype paratype paratype paratype paratype holotype

FL 9.7 12.0 11.9 12.8 12.1 13.4 TBL 12.6 16.7 15.3 15.0 14.0 14.2 AG 20.9 31.0 30.2 28.3 27.0 26.5 HL 13.5 17.7 17.6 17.5 16.8 17.7 HW 8.8 10.4 11.6 11.6 10.8 11.2 HD 5.6 8.8 7.2 7.0 7.0 6.9 ED 3.0 3.1 3.0 3.4 3.1 3.4 EE 3.8 5.0 5.3 5.5 4.4 4.7 ES 7.0 9.0 8.8 8.8 8.4 8.9 EN 5.5 / 6.8 6.6 6.4 6.8 IO 2.8 3.8 3.7 2.6 / 3.5 EL 1.2 1.5 1.3 1.6 1.0 1.6 IN 1.3 / 1.9 1.8 1.7 1.6

Distribution. Cnemaspis sundagekko sp. nov. is known only from the type locality of Pulau Siantan of the Anambas Archipelago, Riau Province, Indonesia (Fig. 4). We expect that it occurs on other nearby islands such as Pulau Matak as well. Mr. F. N. Chasen made collections on Pulau Jimaja to the west and did not report this species. Natural History. Nothing has been reported on the natural history of this species. Being that it is the only species of Cnemaspis thus far known from the Anambas Archipelago we suspect it is a habitat generalist. Comparisons. Within the Southern Seribuat clade, Cnemaspis sundagekko sp. nov. is differentiated from the species of the nigridia group (C. dringi, C. nigridia and C. paripari) by lacking as opposed to having precloacal pores; having fewer paravertebral tubercles (20–25 versus 265–43 collectively); having a single, enlarged, median row of keeled, subcaudal scales in the posterior portion of the tail as opposed to having smooth subcaudals throughout; having caudal tubercles that encircle the tail as opposed to lacking them; and having a greater number of subdigital lamellae on the fourth toe (33–38 versus 26–35 collectively). From the other species of the kendallii group (C. baueri, C. bidongensis, C. kendallii, C. mumpuniae sp. nov., C. pemanggilensis, and C. peninsularis sp. nov.), C. sundagekko sp. nov. differs from C. pemanggilensis by having a much smaller maximum SVL (68.0 mm versus 76.0 mm); fewer paravertebral tubercles (20–25 versus 30–37); a greater number of subdigital lamellae on the fourth toe (33–38 versus 27–34); and white caudal tubercles. From C. mumpuniae sp. nov. it differs in having a larger maximum SVL (68.0 mm versus 60.9 mm) and a greater number of lamellae on the fourth toe (33–38 versus 29–34). From C. baueri, C. sundagekko sp. nov. can be separated on the basis of having a greater number of subdigital lamellae on the fourth toe (33–38 versus 26–32) and a dorsal pattern lacking large black markings as opposed to having such markings. Cnemaspis sundagekko sp. nov. can be differentiated from C. bidongensis by having a larger maximum SVL (68.0 mm versus 58.1 mm); a greater number of subdigital lamellae on the fourth toe (33–38 versus 26–30); and the presence of white caudal tubercles. Cnemaspis sundagekko sp. nov. can be differentiated from C. peninsularis sp. nov. by having a larger maximum SVL (68.0 mm versus 60.0 mm); a greater number of subdigital lamellae on the first toe (33–38 versus 27–33); and posterior one-half of the tail in males being weakly banded as opposed to solid black. Lastly, from C. limi, C. sundagekko sp. nov. is separated by having a much smaller maximum SVL (68.0 mm versus 88.2 mm); fewer paravertebral tubercles (20–25 versus 25–35); a row of ventrolateral caudal tubercles; keeled as opposed to smooth subcaudal scales; and caudal tubercles that do not encircle the tail. Cnemaspis sundagekko sp. nov. can be differentiated from C. sundainsula sp. nov. by having a much smaller maximum SVL (68.0 mm versus 84.5 mm); 11–13 versus 9–11 supralabials; fewer paravertebral tubercles (20–25 versus 26–37); keeled as opposed to smooth subcaudals; caudal tubercles that encircle the tail; and a greater number of subdigital lamellae on the fourth toe (33–38 versus 26–31).

136 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 172 Relationships. Based on distribution, we hypothesize that Cnemaspis sundagekko sp. nov. is part of the Southern Sunda clade and within that clade, it aligns itself with the kendallii group by lacking precloacal pores and having caudal tubercles that encircle the tail (Table 6).

Discussion

Having a well-supported, multilocus phylogeny supporting an unambiguous, morphological taxonomy allows us to entertain other aspects of this group’s evolutionary biology that were intractable prior to this analysis. Much of what can now be done is beyond the scope of this paper, however, below we provide a framework of initial hypotheses and observations that will guide future work—some of which is currently in progress.

Comments on biogeography

Cnemaspis has a unique distribution in that it is generally restricted to the areal, continental fringes of the Sunda Shelf and islands across the southern Sunda Plains (Fig. 1). This, coupled with the phylogenetic relationships and low vagility of its species, has potential in providing insight as to how the cyclical, transitory environmental and geographic history of Sundaland (e.g., Hall 1998, 2001, 2002, 2012; Bird et al. 2005; Outlaw & Voelker 2008; Reddy 2008; Cannon et al. 2009; Woodruff 2010; Cannon 2012; Morley 2012) has influenced the phylogeographic structure of this group (Grismer et al. 2011a). The intractable problem, however, is that key taxa for addressing such issues were most likely distributed across the varied habitats (see Bird et al. 2005) of the Sunda Plains which are now submerged. Thus, any biogeographical hypothesis generated can come only from refugial lineages on the areal fringes of this vast region. The most striking feature concerning the phylogeny of Cnemaspis are the short branch lengths in the deeper regions of the tree between strongly supported clades and species groups that exist in allopatry or parapatry along well-established, biogeographic zones (Figs. 1–4). The exception being the Pattani clade’s distribution within the southern portion of the Northern Sunda clade near the Thai-Malaysian border (Fig. 2). The distribution of the three sequentially basal lineages (the Ca Mau clade from southern Vietnam, the Pattani clade from southernmost Thailand, and the Southern Sunda clade from southern Sundaland) on the opposite sides of the Gulf of Thailand (Fig. 2) would suggest the origin of Cnemaspis was somewhere on the submerged Sunda Plains. The short branch lengths between the Pattani, Northern Sunda, and Southern Sunda clades may indicate a rapid, nearly simultaneous sequence of cladogenic events as inferred from the vertical alignment of their nodes (Figs. 2, 3, 4) followed by radiations into the current species groups. The allopatry of the species groups within the Northern Sunda clade, in most cases, coincides with generally well-studied biogeographical transition zones. The Chao Phraya River basin in southern Thailand that separates the Tenasserim Mountains of western Thailand and Myanmar from the Cardomom Mountains of southern Thailand and Cambodia also separates the chanthaburiensis group from the siamensis group (Figs. 1, 2). The siamensis group is separated from the argus and affinis groups by the Kangar-Pattani Line near the Thai-Malaysian border. This lowland area (< 100 m) runs diagonally from southwest to northeast across the Thai-Malay Peninsula between the Banjaran Titiwangsa Mountains of Peninsular Malaysia in the south and the Sankalakhiri Mountains of Thailand to the north (Fig. 1). It was originally linked to the transition of Malay-type evergreen rain forest into the Thai-Burmese wet seasonal evergreen rain forest (see Van Steenis 1950; Wikramanayake et al. 2000; Woodruff 2003a) and of late has been shown to be an area of faunal exchange in number of groups of lizards, birds, and mammals (Reddy 2008; Woodruff & Turner 2009; Patou et al. 2010; Grismer 2011a). It has been posited, that this region was submerged in the Miocene (24–13 Ma) and again in the Pliocene (5.5–4.5 Ma), creating a 30–100 km wide seaway across the Thai-Malay Peninsula (Woodruff 2003a; however see Parnell 2013). The 17 species of the argus and affinis groups are confined to a relatively small portion of northern Peninsular Malaysia and broadly overlap but are only narrowly parapatric with C. peninsularis sp. nov. of the Southern Sunda clade at the Raub- Betong Suture Zone along the east side of the Banjaran Titiwangsa Mountains (Figs. 2–4). The phylogeographic relationships within the species groups of each clade are reasonably well-resolved. Within the Northern Sunda clade, the chanthaburiensis group occurs along the northern margin of the Gulf of Thailand through the Cardamom Mountains of southern Thailand and Cambodia eastward to the mountainous

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 137 173 regions on the Mekong Delta in southern Vietnam. It is separated from its possible sister lineage (support for this relationship is weak), the siamensis group of the northern Thai-Malay Peninsula by the wide, lowland plains of the Chao Phraya River Basin (Figs. 1, 3) which was flooded during the last Holocene highstand (Woodruff & Turner 2009). Within the chanthaburiensis group, the westernmost sister species C. chanthaburiensis and C. neangthyi are basal to the Vietnamese lineage containing C. aurantiacopes, C. caudanivea, C. tucdupensis, and C. nuicamensis approximately 300 km to the east (Figs. 1–3). The Vietnamese species are restricted to islands or habitat islands associated with the Mekong Delta (Grismer & Ngo 2007). Their well-resolved relationships suggest their phylogenetic structuring happened over a relatively long period of time and is likely tied to the interactions between the complex formation of the Mekong Delta (Thi et al. 2002) and cyclical changes in sea levels (Woodruff & Turner 2009; Woodruff 2010). The siamensis group is composed of a northern (at least two species) and southern (at least three species) lineage from opposite sides of the Isthmus of Kra (Fig. 2)—a well-known biogeographical barrier that separates a large number of species across a wide array of taxonomic groups (e.g., Hughes et al. 2003; Pauwels et al. 2003; Round et al. 2003; Woodruff 2003a,b; De Bruyn et al. 2005; Cattulo et al. 2008; Reddy 2008; Woodruff & Turner 2009; Parnell 2013). Species of the argus and affinis groups occur on both sides of the Banjaran Titiwangsa Mountains—a prominent geographic feature separating the western one-third of Peninsular Malaysia and southern Thailand from the eastern two-thirds along the Raub-Bentong Suture Zone. It is likely this mountain range has played a significant role in the diversification within both these groups being that it separates the basal species of the argus group (C. flavigaster) from the remaining three species (C. karsticola, C. argus, and C. perhentianensis) in the northeast. In the affinis group, this mountain harbors four upland endemics (C. narathiwatensis, C. stongensis sp. nov., C. temiah sp. nov., and C. flavolineata from north to south); restricts six species to the west (C. pseudomcguirei, C. affinis, C. harimau, C. shahruli, C. mcguirei, and C. grismeri) and three species to the east (C. hangus sp. nov., C. selamatkanmerapoh, and C. bayuensis) with no discernable phylogeographic patterns amongst any of them (Figs. 2, 3). The affinis group contains 13 species whose relationships to one another are well-resolved in the ND2 phylogeny and whose topology is represented in Figure 2 although the support for these relationships in the concatenated tree is weak. Cnemaspis pseudomcguirei from the Banjaran Bintang mountains represents the basal species lineage of this group and the sister species C. affinis from Penang Island and C. harimau from Gunung Jerai on the adjacent mainland represents one of the next two basal lineages (Fig. 2). During slightly lower sea levels, these two regions were in contact through a series of low mountains that now exist as a string of islands (Pulau Song, Pulau Bidan and Pulau Telur) lying between them. This same phylogeographic pattern is emerging in other taxa from these areas (Quah et al. in prep.). The other basal lineage of that sister pair contains two clades; one containing the upland species C. narathiwatensis and its lowland sister species C. shahruli to the west and the other clade composing a polytomy of the remaining eight species (Fig. 2). Some of the species relationships within this polytomy, however, are resolved. Cnemaspis mcguirei from the upper regions of the Banjaran Bintang Mountains is the sister species of C. grismeri from the eastern base of the Banjaran Bintang and there is a lineage composed of three small, lowland, karst- dwelling species (C. bayuensis, C. selamatkanmerapoh, and C. hangus sp. nov., the latter two being sister species) from the northeast and an upland, granite-dwelling species C. stongensis sp. nov. that is the sister species of C. bayuensis. The relationships and distribution of the species in the Southern Sunda clade illustrate how cyclical, environmental changes across broad expanses of time can generate sequential, cladogenic events in the same geographic area as evidenced by the position of these events in different places on the tree. The Southern Sunda clade is a polytomous group composed of three major lineages; C. limi from Tioman and Tulai islands, the nigridia group from northern Borneo, and the kendallii group ranging from southern Peninsular Malaysia and Singapore eastward through the Seribuat, Anambas, and Natunas archipelagos to Borneo. Eastern Peninsular Malaysia, the Seribuat and Anambas archipelagos, and the Natuna Archipelago and northwestern Borneo are each composed of separate, Upper Cretaceous-Late Paleocene northwest to southeast oriented arcs of granite plutons (Cobbing et al. 1992; Hutchinson 2009) on which many of the species of this group occur and may account for the origin of the three basal lineages in the Southern Sunda Clade (i.e. C. limi, the nigridia group, and C. sundainsula sp. nov). Speciation events within this clade are likely tied to cyclical changes in sea levels over the last 2.5 million years that would have isolated and reconnected these archipelagos with Peninsular Malaysia and Borneo through

138 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 174 lowland, hilly areas and the intervening archipelagos at least 50 times (Woodruff 2010). Such events could serve as a “species-pump” generating sequential, cladogenic events on the same island arcs which could account for distantly related species occurring on the same islands (e.g., C. limi and C. peninsularis sp. nov. on Pulau Tioman and C. sundainsula sp. nov. and C. mumpuniae on Pulau Natuna Besar). A detailed biogeographic scenario at this point would be premature in that precise phylogenetic information from populations on geographically intermediate granite arcs (C. sundagekko sp. nov. from the Anambas and C. cf. kendallii from the Tambelan archipelagos, respectively) that would be crucial to any scenario are lacking. This issue will be addressed in a forthcoming paper following the acquisition of additional material from these islands.

Comments on diversity

The distribution of the known species of Cnemaspis (Figs. 1, 3, 4) is characterized by large, geographic discontinuities between species across vast regions as well as surprisingly low species diversity in other areas, suggesting that many species remain to be discovered. For example, in Southern Indochina, C. laoensis is currently a geographic outlier to the north, being separated from the nearest population of C. chanthaburiensis in southern Thailand by 600 km (Fig. 1). Given the rocky, mountainous terrain between these two species across southern Laos, Cambodia, and Vietnam we can surmise this hiatus is a collecting artifact as was demonstrated across southern Cambodia and Vietnam with the discovery of C. aurantiacopes, C. nuicamensis, C. caudanivea, C. tucdupensis (Grismer & Ngo 2007) and C. neangthyi (J. Grismer et al. 2010) between C. chanthaburiensis and C. boulengerii filling a hiatus of nearly 600 km. And still a hiatus of nearly 300 km remains between C. neangthyi and C. nuicamensis through suitable lowland, hilly, rocky habitat. The highest diversity of Cnemaspis occurs on the Thai-Malay Peninsula (30 species) with 23 species in Peninsular Malaysia alone. This stands in sharp contrast to the four known species from Borneo (Grismer & Chan 2009), a landmass with nearly four times the surface area and a mountainous terrain with extensive areas of karst and granite formations exceeding those of Peninsular Malaysia. Furthermore, Borneo and Peninsular Malaysia were connected through a land-positive mountainous corridor (the remnants of which are the Natuna, Anambas, and Seribuat archipelagos) as little as 14 thousand years ago (Sathimurthy & Voris 2006.). Despite the number of overlapping and repetitive pocket guides and popular books on the herpetofauna of northern Borneo (see references in Grismer & Chan 2009), only recycled, anecdotal information on Cnemaspis has emerged. We posit that if research on Cnemaspis in Borneo were to commence, its diversity would exceed that of Peninsular Malaysia. We also suspect that Cnemaspis will eventually be found on the large, environmentally diverse island of Sumatra.

Comments on parallel evolution

The various species of Cnemaspis range across a relatively broad array of ecosystems and microhabitats. Some species are strictly lowland forms that do not range above 600 m in elevation, some are upland forms not occurring below 600 m, and others may extend from sea level to 1000 m so long as the appropriate microhabitat is present. Many species are substrate specialists that prefer only one type of rock surface (limestone or granite), others are vegetational specialists or habitat generalists, and one species, C. chanthaburiensis, is strictly terrestrial and can be found beneath and within logs and beneath rocks. Across all these environmental gradients, many species are nocturnal, others are diurnal, and some are both. The phylogenetic hypothesis of Cnemaspis indicates substrate preferences, activity periods, elevational zonation, and the presence of ocelli has evolved in parallel multiple times (Fig. 5). We note here that the most common combination of traits for Cnemaspis is to be a diurnal, granite- dwelling, lowland species lacking ocelli (19 of 44 species examined for which molecular data were available; Table 16). Ancestral state reconstructions based on maximum likelihood analyses will be used to test hypotheses concerning the evolutionary origins and trends among these different parameters are currently in progress (Wood & Grismer in prep.).

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 139 175 TABLE 16. Activity period, substrate preference (microhabitat), elevation (upland > 600 m), and presence or absence of ocelli (eyespots) in the species of Cnemaspis for which we had sufficient data or observations. Activity period Substrate preference Elevation Ocelli location nocturnal diurnal limestone granite vegetation terrestrial upland lowland shoulders head neck

Ca Mau clade boulenegerii X X X psychedelica X X X

Pattani clade monachorum X X X biocellata X X X X X X niyomwanae X X X kumpoli X X X X

Northern Sunda clade

chanthaburiensis group chanthaburiensis X X X X neangthyi X X X aurantiacopes X X X caudanivea X X X nuicamensis X X X tucdupensis X X X

siamensis group siamensis X X X huaseesom X X X chanardi X X X X omari sp. nov. X X X roticanai X X X X kamolnorranathi X X X X X punctatonuchalis X X X X X vandeventeri X X X

argus group flavigaster X X X X argus X X X X karsticola X X X perhentianensis X X X

affinis group pseudomcguirei X X X X harimau X X X X X affinis X X X X shahruli X X X X X temiah sp. nov. X X X flavolineata X X X hangus sp. nov. X X X selamatkanmerapoh X X X stongensis sp. nov. X X X X bayuensis X X X mcguirei X X X X grismeri X X X X narathiwatensis X X X X

Southern Sunda clade limi X X X X

nigridia group nigridia X X X paripari X X X

kendallii group kendallii X X X X X sundainsula sp. nov. X X X pemanggilensis X X X baueri X X X mumpuniae sp. nov. X X X X bidongensis X X X X peninsularis sp. nov. X X X X X X

140 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 176 Comments on genetic divergence with Cnemaspsis

The genetic divergence based on ND2 seen among species of Cnemaspis greatly exceeds that seen among species within other gekkotan genera such as Hemiphyllodactylus, Phyllopezus, Pseudogekko, Ptychozoon, Cyrtodactylus.

Comments on integrative taxonomy

A general assumption concerning integrative taxonomic analyses is that currently recognized, widespread species are composed of morphologically cryptic species whose detection is possible only through the use of molecular analyses (see Grismer et al. 2013b for a discussion). On the contrary, the majority of integrative analyses result in demonstrating that diagnostic, morphological characters were present but overlooked by the last author(s) to revise the group and their subsequent discovery was simply prompted by the results of a molecular analysis (see references in Grismer et al. 2013b). Prior to 2003, Cnemaspis was known from 10 species. Between 2003 and this study, we and associated authors described 34 of the then 44 known species on the basis of morphology and color pattern alone (Das & Grismer 2003; Grismer & Das 2006; Grismer & Ngo 2007; Chan & Grismer 2008; Grismer & Chan 2008, 2009, 2010; Grismer et al. 2008a,b, 2009, 2010a,b,c; Grismer 2009; Chan et al. 2010; J. Grismer et al. 2010; Wood et al. 2013). The analysis herein indicated the only error made was that a geographically outlying specimen from southern Thailand (included as part of the type series of C. chanardi) and continental populations of C. roticanai actually constituted a new species, C. omari sp. nov., which turned out to be the sister species of C. roticanai sensu stricto. This mistake was made because Grismer et al. (2010a) and Grismer & Chan (2010) did not have all the material available on hand to compare. The molecular analysis indicated these were different taxa (Fig. 2), prompting a reexamination herein by comparing all the material together and in so doing, we discovered additional, diagnostic, morphological characters. Thus, integrative taxonomic analyses are not necessarily revealing cryptic species but rather highlighting less than efficient morphological analyses.

Acknowledgements

We wish to thank Alexandra Sumaril, Jacob Chan, Maria Kuss, Mathew Murdoch, Heather Heinz, Ariel Loreado, Michael Cota, and Rick Gregory for field assistance. We thank the BMNH (E. N. Arnold and C. J. McCarthy), CAS (A. E. Leviton and J. Vindum), FMNH (H. K. Voris, R. F. Inger, and A. Resetar), MCZ (J. E. Cadle and J. P. O. Rosado), MSNG (R. Poggi and G. Doria), UF (D. Auth and F. W. King), USDZ (ZRC in Leviton et al., 1985); K. K. P. Lim, P. K. L. Ng, H. H. Tan, and C. M. Yang; USNM (R. I. Crombie, W. R. Heyer, and G. R. Zug), ZMA (A.Groenveld and L. van Tuijl), G. L. Lenglet and S. Brauxof the Institut Royal des Sciences naturelles de Belgique; and ZSI (J. R. B. Alfred, S. K. Chanda, B. Dattagupta and N. C. Gayen) for permitting us to examine material under their care. Lastly we are most thankful to Tanya Chan-ard of the Thailand Natural History Museum, National Science Museum (THNHM), Kumthorn Thirakhupt of the Chulalongkorn University Museum of Zoology (CUMZ), Dr. Sansareeya Wangkulangkul of the Prince of Songkhla University Zoological Collection (PSUZC), Wut Taksinthum of the Zoological Museum of Kasetsart University (ZMKU), Apirat Taokratok of the Nakhonratchasima Zoo Museum, ZPO (KZM) whose generous and expeditious loan of material made this project possible. We thank P. David (MNHN) for useful information and H. Bringsøe who provided a picture of C. vandeventeri. We thank Dr. Jack W. Sites Jr. for partial founding of the molecular analyses. AR thank to Local Government of Natuna especially Dinas Bunhut and BPLHD for funded research in Natuna. This research was supported in part by a grant to LLG from the College of Arts and Sciences, La Sierra University, Riverside, California and by Ministry of Higher Education Grant to Shahrul Anuar.

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146 · Zootaxa 3880 (1) © 2014 Magnolia Press GRISMER ET AL. 182 Woodruff, D.S. (2010) Biogeography and conservation in Southeast Asia: how 2.7 million years of repeated environmental fluctuations affect today's patterns and the future of the remaining refugial-phase biodiversity. Biodiversity Conservation, 19, 919–941. http://dx.doi.org/10.1007/s10531-010-9783-3 Woodruff, D.S. & Turner, L.M. (2009) The Indochinese–Sundaic zoogeographic transition: a description of terrestrial mammal species distributions. Journal of Biogeography, 36, 803–821. http://dx.doi.org/10.1111/j.1365-2699.2008.02071.x Youmans, T.M., Grismer, J.L., Escobar III, R.A., Johnson, R. & Grismer, L.L. (2002) First report on the herpetofauna of Pulau Pemanggil, Johor, West Malaysia. Hamadryad, 27, 148–149.

MONOGRAPH OF SOUTHEAST ASIAN CNEMASPIS Zootaxa 3880 (1) © 2014 Magnolia Press · 147 183 Chapter 4: Three new karst-dwelling Cnemaspis Strauch, 1887 (Squamata; Gekkonidae) from Peninsular Thailand and the phylogenetic placement of C. punctatonuchalis and C. vandeventeri

184 Three
new
karst-dwelling
Cnemaspis
Strauch,
1887
(Squamata;
GekkoniEBF)
from
Peninsular
Thailand
and
the
phylogenetic
placement
of C.
punctatonuchalis
and
C.
vandeventeri

Perry Lee Wood Jr1, L. Lee Grismer2, Anchalee Aowphol3, César A. Aguilar1, Micheal Cota4,5, Marta S. Grismer2, Matthew L. Murdoch2 and Jack W. Sites Jr1 1 Department of Biology and Bean Life Science Museum, Brigham Young University, Provo, UT, United States 2 Department of Biology, La Sierra University, Riverside, CA, United States 3 Faculty of Science, Department of Zoology, Kasetsart University, Chatuchak, Bangkok, Thailand 4 Natural History Museum, National Science Museum, Thailand, Technopolis, Khlong 5, Khlong Luang, Pathum Thani, Thailand 5 Suan Sunandha Rajabhat University, Institute for Research and Development, Dusit, Bangkok, Thailand

ABSTRACT Three new species of Rock Geckos Cnemaspis lineogularis sp. nov., C. phangngaensis sp. nov., and C. thachanaensis sp. nov. of the chanthaburiensis and siamensis groups are described from the Thai portion of the Thai-Malay Peninsula. These new species are distinguished from all other species in their two respective groups based on a unique combination of morphological characteristics, which is further supported by mitochondrial DNA (mtDNA) from the NADH dehydrogenase subunit 2 gene (ND2). Cnemaspis lineogularis sp. nov. is differentiated from all other species in the chanthaburiensis group by having a smaller maximum SVL 38 mm, 13 paravertebral tubercles, enlarged femoral scales, no caudal bands, and a 19.5–23.0% pairwise sequence divergence (ND2). Cnemaspis phangngaensis sp. nov. is differentiated from all other species in the siamensis group by having the unique combination of 10 infralabial scales, four continuous pore-bearing precloacal scales, paravertebral tubercles linearly Submitted 7 September 2016 Accepted 8 December 2016 arranged, lacking tubercles on the lower flanks, having ventrolateral caudal tubercles Published 24 January 2017 anteriorly present, caudal tubercles restricted to a single paraveterbral row on each Corresponding author side, a single median row of keeled subcaudals, and a 8.8–25.2% pairwise sequence Perry Lee Wood Jr, [email protected] divergence (ND2). Cnemaspis thachanaensis sp. nov. is distinguished from all other Academic editor species in the siamensis group by having 10 or 11 supralabial scales 9–11 infralabial Keith Crandall scales, paravertebral tubercles linearly arranged, ventrolateral caudal tubercles ante- Additional Information and riorly, caudal tubercles restricted to a single paravertebral row on each side, a single Declarations can be found on median row of keeled subcaudal scales, lacking a single enlarged subcaudal scale row, page 40 lacking postcloaclal tubercles in males, the presence of an enlarged submetatarsal scale DOI 10.7717/peerj.2884 at the base if the 1st toe, and a 13.4–28.8% pairwise sequence divergence (ND2). The new phylogenetic analyses place C. punctatonuchalis and C. vandeventeri in the Copyright 2017 Wood Jr et al. siamensis group with C. punctatonuchalis as the sister species to C. huaseesom and C. vandeventeri as the sister species to C. siamensis, corroborating previous hypotheses Distributed under based on morphology. The discovery of three new karst-dwelling endemics brings the Creative Commons CC-BY 4.0 total number of nominal Thai Cnemaspis species to 15 and underscores the need for OPEN ACCESS

How to cite this article Wood Jr et al. (2017), Three new karst-dwelling Cnemaspis Strauch, 1887 (Squamata; Gekkoniade) from Peninsu- lar Thailand and the phylogenetic placement of C. punctatonuchalis and C. vandeventeri. PeerJ 5:e2884; DOI 10.7717/peerj.2884 185 continued field research in poorly known areas of the Thai-Malay Peninsula, especially those that are threatened and often overlooked as biodiversity hot spots.

Subjects Genetics, Taxonomy, Zoology Keywords Limestone forests, chanthaburiensis group, siamensis group, Malay peninsula

INTRODUCTION The Thai-Malay Peninsula is a long (1,127 km) and narrow (maximum width 322 km) appendix of mainland Asia extending from Indochina in the north to its southern terminus in Singapore. The Thai-Malay Peninsula is comprised of the southern portion of Myanmar, the southwestern section of Thailand, West Malaysia, and Singapore. This region is both geologically and climatically complex and has been influenced by a number of factors. The environmental complexity of this region has helped to form two prominent biogeographic barriers, the Isthmus of Kra and the Kangar-pattani line. These biogeographic barriers serve as pivotal crossroads for faunal exchange between the Indochinese and Sundaic biota (e.g., Raes et al., 2014; De Bruyn et al., 2013; Parnell, 2013; Patou et al., 2009; Woodruff & Turner, 2009; Gorog, Sinaga & Engstrom, 2004; Pauwels et al., 2003; Hughes, Round & Woodruff, 2003; Woodruff, 2003; Grismer et al., 2014d; Grismer, 2011). One feature that is often over- looked in terms of biodiversity are the myriad of limestone forests and karst formations dispersed throughout the Malay Peninsula. Karstic regions have been referred to as ‘‘arks’’ or biodiversity reservoirs that can be used as stock for repopulating degraded environments during ecosystem reassembly (Schilthuizen, 2004). In addition to serving as arks, karst formations have been known to provide natural laboratories for biogeographic, evolutionary, ecological, and taxonomic research (e.g., Ng, 1991; Grismer et al., 2014c; Grismer et al., 2014b; Schilthuizen et al., 2005; Schilthuizen et al., 1999; Kiew, 1991). From chemical and mechanical weathering karst formations have been molded into a unique suite of microhabitats in which a number of species have become adapted (e.g., Vermeulen & Whitten, 1999; Komo, 1998a; Komo, 1998b; Tija, 1998). To date there has been a fair amount of research conducted on the flora of karst formations and their surrounding limestone forests, resulting in a high estimate of endemic species (Kiew, 1998; Clements et al., 2006; Chin, 1977 and references therein). In addition to the high level of floral endemism there are also high levels of invertebrate endemism associated with karst formations (e.g., Holloway, 1986; Vermeulen & Whitten, 1999). Although these areas harbor a high degree of endemism for invertebrates and plant species they are generally not considered to hold large numbers of endemic terrestrial vertebrates (i.e., Jenkins et al., 2005; Alström et al., 2010; Woxvold, Duckworth & Timmins, 2009), because most vertebrates have high dispersal capabilities. There are only a few mammals and birds that are thought to be restricted to karst formations (e.g., Latinne et al., 2011; Clements et al., 2006). In contrast, recent taxonomic work in Peninsular Malaysia has uncovered an impressive amount of new microendemic karst-dwelling species of reptiles, including 14 new lizards (Grismer et al., 2008a; Grismer et al., 2008b; Grismer

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 2/46 186 et al., 2009; Grismer et al., 2013b; Grismer et al., 2012; Grismer et al., 2014e; Grismer et al., 2014c; Grismer et al., 2013a; Grismer et al., 2016a; Wood Jr et al., 2013) and two new snakes (Grismer et al., 2014b; E Quah, 2017, unpublished data). However, these surveys only covered a small portion of the limestone forests and karst formations of Peninsular Malaysia continue northward up the entire Thai-Malay Peninsula into central Thailand and eastern Myanmar. Dispersed throughout Peninsular Thailand are hundreds of unexplored isolated karst formations. From the limited number of surveys that have been conducted, a few gekkotan species have been identified and described (e.g., Grismer et al., 2012; Grismer et al., 2015; Pauwels et al., 2013; Grismer et al., 2012; Ellis & Pauwels, 2012). A major focus of these surveys in the last two decades has lead to the discovery of at least 15 new micro-endemic karst-dwelling Bent-toed Geckos in the genus Cyrtodactylus (e.g., Ellis & Pauwels, 2012; Pauwels et al., 2013; Grismer et al., 2016a; Sacha, 2015, and references therein). The genus Cyrtodactylus is a monophyletic assemblage of 224 species (Uetz, Freed & Ho≤ek, 2016) that have a broad range throughout Asia to the western Pacific (Wood Jr et al., 2012). This diverse group of nocturnal geckos occupies a broad suite of microhabitats (eg. karst and granite caves, granite boulders, leaf-litter, limestone forests etc.) with a number of specialists in distantly related clades. One group of convergent specialists that has recently received attention are the karst and cave dwelling ecomoprhs in the condorensis and intermedius species complexes (Grismer et al., 2015, and references therein). These granite and karst-dwelling Cyrtodactylus are often found syntopically with the Rock Geckos in genus Cnemaspis and can pose as a potential competitor. The Rock geckos of the genus Cnemaspis comprise a clade of 55 described species that are widespread throughout the Sunda Shelf, with a majority of the species being from the Thai-Malay Peninsula and their adjacent islands. Most Cnemaspis are diurnal, cryptically colored, scansorial species, however some species such as C. psychedelica Grismer, Ngo & Grismer 2010 (Fig. 8 in Grismer et al., 2014d) are brightly colored and a number of species dispersed throughout the phylogenetic tree are nocturnal with multiple independent transitions (Fig. 5 in Grismer et al., 2014d). Like Cyrtodactylus, Cnemaspis are also found on a variety of substrates (e.g., granite, karst, vegetation, terrestrial and various combinations of these). Often when there are micro-endemic karst-dwelling Cyrtodactylus (nocturnal) there is usually an endemic diurnal karst-dwelling Cnemaspis occupying the same niche during different activity periods (e.g., Grismer et al., 2016a; Grismer et al., 2012; Grismer et al., 2014e; Grismer et al., 2016b). From this arises a number of interesting questions about niche partitioning, behavior, and potential competition of Cyrtodactylus and Cnemaspis, however this is not the focus of this paper. Recent surveys in Phangnga, Tha Chana, Thap Sakae, Prachuap Khiri Khan, and Tham Sonk hill during the month of September 2016 resulted in the collection of Cnemaspis punctatonuchalis, C. vandeventeri and three undescribed species of Cnemaspis that co-occur with some of the aforementioned species of karst endemic Cyrtodactlyus. These new populations can be placed in the genus Cnemaspis based on having broad, flattened heads; large somewhat forward and upwardly directed eyes with round pupils and no eyelids; flattened bodies; long, widely splayed limbs with long, inflected digits; and no femoral pores. Here we present morphological and color

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 3/46 187 pattern data as evidence, for delimitation of these three new species of Cnemaspis, bolstered by mtDNA genetic data, and present the phylogenetic placement of C. punctatonuchalis and C. vandeventeri.

MATERIALS AND METHODS Taxon sampling We obtained 203 samples of Cnemaspis and outgroups from Grismer et al. (2014d). In combination with this dataset we added 14 new samples including two species of Thai Cnemaspis (C. punctatonuchalis and C. vandeventeri) that have never been sequenced, along with three undescribed species of Cnemaspis from peninsular Thailand (Fig. 1, Table S1). Brigham Young University’s Institutional Animal Care and Use Committee (IACUC) has approved the animal use protocol for this study (protocol # 160401). The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZN), and hence the new names contained in the electronic version are effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub:987831FC- F4BA-4409-A43C-9929F913E9F9. The online version of this work is archived and available from the following digital repositorie: PubMed Central.

Molecular and phylogenetic analyses Genomic DNA was isolated from liver or muscle tissues stored in 95% ethanol using the animal tissue protocol in the Qiagen DNeasyTM tissue kit (Valencia, CA, USA). The mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and the flanking tRNAs ( 1,335 bp) was amplified using a double-stranded Polymerase Chain Reaction (PCR) ⇠ under the following conditions: 1.0 µl( 10–33 µg) genomic DNA, 1.0 µl (10 µM) ⇠ forward primer L4437b (50-AAGCAGTTGGGCCCATACC-30), 1.0 µl (10 µM) reverse primer H5934 (50-AGRGTGCCAATGTCTTTGTGRTT-30), 1.0 µl deoxynucleotide pairs (1.5 µM), 2.0 µl 5x buffer (1.5 µM), 1.0 MgCl 10x buffer (1.5 µM), 0.18 µl Promega Taq polymerase (5 u/µl), and 7.5 µl H2O, primers are from Macey et al. (1997). All PCR reactions were executed in an Eppendorf Mastercycler gradient theromocycler under the following conditions: initial denaturation at 95 C for 2 min, followed by a second denaturation at 95 C for 35 s, annealing at 52 C for 35 s, followed by a cycle extension at 72 C for 35s, for 33 cycles. All PCR products were visualized on a 1% agarose gel electrophoresis. Successful targeted PCR products were vacuum purified using MANU 30 PCR Millipore plates and purified products were resuspended in DNA grade water. Purified PCR products were sequenced using the PCR primers from above and sequencing primers CyrtintF1 (50-TAGCCYTCTCYTCYATYGCCC-30) and CyrtintR1 (50-ATTGTKAGDGTRGCYAGGSTKGG-30) from (Siler et al., 2010) on the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 4/46 188 Thailand Laos

14

Cambodia

Vietnam

Myanmar

9 siamensis chanthaburiensis group group

vandeventeri aurantiacopes chanardi caudanivea huaseesom chanthaburiensis kamolnorranathi* neangthyi omari lineogularis sp. nov. phangngaensis sp. nov. nuicamensis punctatonuchalis tucdupensis 0 50 km roticanai siamensis 99 Malaysia 104 thachanaensis sp. nov. 109

Figure 1 Distribution of the species of Cnemaspis in the chanthaburiensis and siamensis groups. Stars indicate type localities, colored dots represent additional localities for the respective species, and the col- ored outlines correspond to colored clades in Fig. 2. The asterisk (*) identifies a species not included in the molecular analysis and is hypothesized to be in the siamensis group based on its distribution (Grismer et al., 2014d).

Cycle sequencing reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequenced on an ABI 3730xl DNA Analyzer at the BYU DNA Sequencing Center (DNASC). All new sequences produced from this study are deposited in GenBank under the following accession numbers KY091231–KY091244 (Table S1). All sequences were edited and aligned in Geneious v6.1.8 (Kearse et al., 2012), alignment was constructed using the Muscle plugin (Edgar, 2004). Mesquite v3.02 (Maddison & Maddison, 2015) was used to check for stop codons and to ensure the correct amino acid read frame. For estimating the phylogenetic relationships we used both partitioned Maximum Likelihood (ML) and partitioned Bayesian Inference (BI) methods. The ND2 gene was partitioned by codon position and the tRNAs were treated as a single partition for both the ML and BI analyses. All models of molecular evolution were estimated in ModelTest v3.7 (Posada & Crandall, 1998), using the Bayesian Information Criterion (BIC). The best fit

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 5/46 189 Table 1 Models of molecular evolution used for the ML and BI analyses.

Gene Model selected Model applied for ML Model applied for BI ND2 1st pos GTR I 0 GTR I 0 GTR I 0 + + + + + + 2nd pos GTR I 0 GTR I 0 GTR I 0 + + + + + + 3rd pos GTR I 0 GTR I 0 GTR I 0 + + + + + + tRNAs TrN I 0 GTR I 0 HKY I 0 + + + + + +

models of evolution are in presented in Table 1. The partitioned ML analyses was performed using RAxML HPC v7.5.4 (Stamatakis, 2006), 1,000 bootstrap pseudoreplicates via therapid hill-climbing algorithm (Stamatakis, Hoover & Rougemont, 2008) with 200 searches for the best tree. The Bayesian analysis was carried out in MrBayes v3.2 (Huelsenbeck et al., 2001; Ronquist et al., 2012) using the default priors. Two simultaneous runs were performed with eight chains per run, seven hot and one cold following default priors. The analysis was run for 2 x 106 generations and sampled every 1000 generations from the Markov Chain Monte Carlo (MCMC). The analysis was halted after the average standard deviation split frequency was below 0.01 and we assumed convergence. We conservatively discarded the first 25% of the trees as burnin and constructed a consensus tree using the sumt command in MrBayes. Nodes having bootstrap support values (BS) greater than 70 and posterior probabilities (PP) above 0.95 were considered well supported (Huelsenbeck et al., 2001; Wilcox et al., 2002). We calculated uncorrected percent sequence divergences for ND2 in Mega v6.06 (Tamura et al., 2013).

Morphological analyses Morphological and color pattern characteristics follow Grismer et al. (2014d): color pattern characters were taken from digital images of living specimens cataloged in the La Sierra University Digital Photo Collection (LSUDPC) and from living specimens in the field. The following measurements on the type series were taken by PLWJ with a electronic digital caliper to the nearest 0.1 mm, under a Lica WILD M10 dissecting microscope on the left side of the body where appropriate: snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90 to the inflection of the flexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90 to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eyeball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eyeball; eye to snout distance (ES), measured from anteriormost margin of the eyeball to the tip of

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 6/46 190 snout; eye to nostril distance (EN), measured from the anteriormost margin of the eyeball to the posterior margin of the external nares; inner orbital distance (IO), the width of the frontal bone at the level of the anterior edges of the orbit; ear length (EL), the greatest vertical distance of the ear opening; and internarial distance (IN), measured between the medial margins of the nares across the rostrum. Additional character states evaluated were numbers of supralabial and infralabial scales counted from below the middle of the orbit to the rostral and mental scales, respectively; the texture of the scales on the anterior margin of the forearm; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column (where applicable); the presence or absence of a row of enlarged, widely spaced, tubercles along the ventrolateral edge of the body (flank) between the limb insertions; the general size (i.e., strong, moderate, weak) and arrangement (i.e., random or linear) of the dorsal body tubercles; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the distribution of transverse and granular subdigital lamellae on the fourth toe; the total number of precloacal pores, their orientation and shape; the number of precloacal scales lacking pores separating the left and right series of pore-bearing precloacal scales; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales, subtibial scales, and submetatarsal scales beneath the first metatarsal; and the number of postcloacal tubercles on each side of the tail base. Longitudinal rows of caudal tubercles on the non-regenerated portion of the tail are quite variable between species and useful in differentiating several taxa. Up to five pairs of the following rows may be present in varying combinations: paravertebral row—the dorsal row adjacent to the middorsal, caudal furrow; dorsolateral row—the row between the paravertebral row and the lateral, caudal furrow on the dorsolateral margin of the tail; lateral row—the row immediately below the lateral, caudal furrow; and ventrolateral row—the row below the lateral row on the ventrolateral margin of the tail below the lateral caudal furrow. When present, this row is usually restricted to the anterior 25% (or less) of the tail. Occasionally there may be a row of tubercles within the lateral, caudal furrow.

RESULTS The phylogenetic analyses place both C. punctatonuchalis and C. vandeventeri in the siamensis group (Fig. 2). Cnemaspis punctatonuchalis is strongly recovered for the ML analysis (100 BS) but not the BI (0.87 PP) as the sister species to C. huaseesom. Cnemaspis vandeventeri is strongly supported (100 BS and 0.99 PP) as the sister lineage to C. siamensis. Phylogenetic analyses of the three new populations sampled from Prachuap Khiri Khan, Phangnga, and Tha Chana represent well-supported independent lineages (100 BS, 1.0 PP; 100 BS, 1.0 PP; 100 BS, 1.0 PP, respectively). The samples from Wat Khao Daeng are well-supported (100 BS, 1.0 PP) as the sister lineage to the chanthaburiensis group (Fig. 2A) and demonstrate a 19.5–23% mtDNA pairwise sequence divergence from all of the other species in this group (Table 2). Both the Phangnga and the Tha Chana populations are nested within the siamensis group (Fig. 2B). The Phangnga population is well-supported for ML (99 BS) but lacks support from the BI (0.56 PP) as the sister lineage to a clade

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 7/46 191 C. lineogularis sp. nov. BYU 62535 Wat Khao Daeng, TH A. 100/1.0 C. lineogularis sp. nov. BYU 62536 Wat Khao Daeng, TH C. lineogularis sp. nov. ZMKU R 00728 Wat Khao Daeng, TH C. aurantiacopes LSUHC 8612 Hon Dat, VT 100/1.0 C. aurantiacopes LSUHC 8611 Hon Dat, VT 100/1.0 9 1/0.82 C. aurantiacopes LSUHC 8610 Hon Dat, VT C. chanthaburiensis LSUHC 9338 Dalai, CM 100/1.0 C. neangthyi LSUHC 8517 O’Lakmeas, CM 100/1.0 C. neangthyi LSUHC 8516 O’Lakmeas, CM C. neangthyi LSUHC 8515 O’Lakmeas, CM

9 2/1.0 C. neangthyi LSUHC 8478 O’lakmeas, CM C. caudanivea LSUHC 8582 Hon Tre, VT 100/1.0 C. caudanivea LSUHC 8578 Hon Tre, VT C. caudanivea LSUHC 8577 Hon Tre, VT C. tucdupensis LSUHC 8632 Tuc Dup, VT 100/1.0 9 7/0.99 C. tucdupensis LSUHC 8631 Tuc Dup, VT C. tucdupensis LSUHC 8634 Tuc Dup, VT 9 8/1.0 C. tucdupensis LSUHC 8633 Giang, VT C. nuicamensis LSUHC 8648 Nui Cam, VT 100/1.0 C. nuicamensis LSUHC 8646 Nui Cam, VT C. nuicamensis LSUHC 8649 Nui Cam, VT C. nuicamensis LSUHC 8647 Nui Cam, VT B. C. chanardi LSUHC 9567 Nakhon, TH 100/1.0 100/1.0 C. phangngaensis sp. nov. BYU 62538 Phangnga, TH C. phangngaensis sp. nov. BYU 62537 Phangnga, TH 100/1.0 C. omari LSUHC 9565 Satun, TH 99/0.56 C. omari LSUHC 9978 Perlis, WM Cnemaspis 100/0.63 C. roticani LSUHC 11815 Pulau langkawi, WM C. roticani LSUHC 9430 Pulau Langkawi, WM 100/1.0 100/1.0 C. roticani LSUHC 9439 Pulau Langkawi, WM C. roticani LSUHC 9431 Pulau Langkawi, WM 100/1.0 C. punctatonuchalis BYU 62540 Thap sakae, TH 100/0.87 C. punctatonuchalis BYU 62539 Thap sakae, TH C. huaseesom LSUHC 9457 Sai Yok, TH 100/1.0 C. huaseesom LSUHC 9458 Sai Yok, TH 100/1.0 C. huaseesom LSUHC 9455 Sai Yok, TH C. vandeventeri BYU 62541 Naka, TH 100/0.99 100/1.0 C. siamensis LSUHC 9485 Chumpon, TH 100/1.0 C. siamensis LSUHC 9474 Chumpon, TH C. thachanaensis sp. nov. BYU 62543 Tha chana, TH 100/1.0 C. thachanaensis sp. nov. ZMKU R 00730 Tha chana, TH C. thachanaensis sp. nov. BYU 62544 Tha chana, TH C. thachanaensis sp. nov. ZMKU R 00729 Tha chana, TH C. thachanaensis sp. nov. ZMKU R 00731 Tha chana, TH 0.01 C. thachanaensis sp. nov. BYU 62542 Tha chana, TH

Figure 2 Phylogenetic relationships of the chanthaburiensis (A) and the siamensis (B) groups. Right, Maximum Likelihood tree from RAxML ( ln L 60818.390304) for all species of Cnemaspis and outgroups with bootstrap support values (BS) and Bayesian posterior probabilities (PP), respectively. Country abbre- viations for the tip labels are as follows: CM, Cambodia; TH, Thailand; WM, West Malaysia; VT, Vietnam. All new species are highlighted with grey boxes and the genus Cnemaspis is highlighted using a box with dashed lines in the main tree.

composed of C. omari and C. roticani and demonstrate a 8.8–25.2% mtDNA pairwise sequence divergence from all of the other species in the siamensis group (Table 3). The population from Tha Chana forms a well-support lineage (100 BS and 1.0 PP) and is strongly (100 BS and 1.0 PP) placed as the sister lineage to a clade composed of C. siamensis and C. vandeventeri and bares a 13.4–28% mtDNA pairwise sequence divergence form all of the other species in the siamensis group (Table 3). Given that these new populations form well-supported independent lineages (Figs. 2A and 2B) coupled with high genetic distances and a unique set of morphological and color pattern characteristics that separate them from all members of their respective groups, we describe these three populations below as new species.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 8/46 192 Table 2 Pairwise uncorrected p-distances based on 1,335 bp of ND2 and associated tRNAs calculated in MEGA v6.06 (Tamura et al., 2013) within the chanthaburiensis group. Within species distances are presented in bold text and between species distances are presented below the diag- onal. C. lineogularis C. aurantiacopes C. caudanivea C. chanthaburiensis C. neangthyi C. nuicamensis C. tucdupensis sp. nov. C. lineogularis 0.003 sp. nov. C. aurantiacopes 0.200 0.011 C. caudanivea 0.195 0.136 0.002 C. chanthaburiensis 0.217 0.161 0.167 – C. neangthyi 0.230 0.161 0.169 0.156 0.001 C. nuicamensis 0.207 0.158 0.160 0.175 0.179 0.002 C. tucdupensis 0.206 0.149 0.140 0.171 0.166 0.140 0.006

Table 3 Pairwise uncorrected p-distances based on 1,335 bp of ND2 and associated tRNAs calculated in MEGA v6.06 (Tamura et al., 2013) within the siamensis group. Within species distances are presented in bold text and between species distances are presented below the diagonal. C. thachanaensis sp. nov. C. phangngaensis sp. nov. C. omari C. chanardi C. punctatonuchalis C. vandeventeri C. huaseesom C. roticani C. siamensis C. thachanaensis sp. nov. 0.003 C. phangngaensis sp. nov. 0.260 0.002 C. omari 0.282 0.112 0.042 C. chanardi 0.243 0.114 0.118 – C. punctatonuchalis 0.211 0.250 0.264 0.255 – C. vandeventeri 0.144 0.252 0.266 0.240 0.211 – C. huaseesom 0.208 0.237 0.282 0.262 0.170 0.201 0.004 C. roticani 0.275 0.088 0.090 0.117 0.256 0.268 0.274 0.002 C. siamensis 0.134 0.250 0.278 0.255 0.194 0.123 0.194 0.281 –

Systematics Cnemaspis lineogularis sp. nov. urn:lsid:zoobank.org:act:8E3B21A4-93BF-4D08-B8D1-0A3EEF6BE44F Common name: Striped Throated Rock Gecko (Figs. 3–5)

Holotype. BYU 62535 adult male, collected near Wat Khao Daeng, Kui Buri, Prachuap Khiri Khan, Thiland (12.134620N, 99.961078E; 12 m a.s.l.), 31 July 2016, by PLW, LLG, CA, MC, MSG, MLM. Paratopotypes. BYU 62536 adult male and ZMKU R 00728 adult female paratypes bear the same collection and data as the holotype. Diagnosis. Cnemaspis lineogularis is distinguished from all other species of Cnemaspis in the chanthaburiensis group by the combination of the following morphological and

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 9/46 193 Figure 3 (A) male holotype BYU 62535 and (B) female paratype ZMKU R 00728 of Cnemaspis lineogu- laris sp. nov.

color pattern characters: maximum SVL 38 mm; nine supralabials; eight infralabials; ventral scales smooth; no precloacal pores; 13 paravertebral tubercles linearly arranged; no tubercles on the lower flanks; lateral caudal furrows present; no caudal tubercles in the lateral furrows; ventrolateral caudal tubercles anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a single median row of enlarged smooth scales; lateral caudal tubercle row absent; shield-like subtibial scales absent; one post cloacal tubercle in males; no enlarged femoral or submetatarsal scales; enlarged femoral scales; subtibials smooth; 27–29 subdigital fourth toe lamellae; sexually dimorphic for dorsal color pattern; gular

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 10/46 194 Figure 4 Ventral coloration and sexual dichromatism in the type series of Cnemaspis lineogularis sp. nov. (A) adult male holotype BYU 62535, (B) adult male paratype BYU 62536, (C) adult female paratype ZMKU R 00728.

region yellow-orange, thick, black lineate markings in males, absent in females; subcaudal region whitish (Tables 4–6). Description of the holotype. Adult male; SVL 38 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.25), somewhat narrow (HW/SVL 0.16), flattened (HD/HL 0.38), head distinct from neck; snout moderate (ES/HL 0.52), snout slightly concave in lateral view; postnasal region concave medially; scales on rostrum smooth becoming keeled posteriorly, raised, larger than conical scales on occiput; weak to absent supra ocular ridges; frontalrostralis sulcus deep; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.26); extra-brillar, fringe scales largest anteriorly; pupil round; ear opening more round than oval; rostral slightly concave, dorsal 80% divided by longitudinal median

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 11/46 195 Figure 5 Dorsal view of the type series of Cnemaspis lineogularis sp. nov. (A) adult male holotype BYU 62535, (B) adult male paratype BYU 62536, (C) adult female paratype ZMKU R 00728.

groove; rostral bordered posteriorly by supra nasals and one small azygous scale and laterally by first supralabials; 9,9 (R,L) slightly raised supralabials decreasing in size posteriorly; 8,8 (R,L) infralabials decreasing in size posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered by small postnasal scales; mental large, triangular, concave, bordered posteriorly by three postmentals; gular and throat scales raised, smooth, small and round. Body slender, elongate (AG/SVL 0.46); small, keeled, dorsal scales equal in size throughout body, intermixed with several large, multicarinate conical tubercles more or less randomly arranged; tubercles extend from the occiput to base of the tail; no tubercles on flanks; pectoral and abdominal scales smooth, not larger posteriorly; abdominal scales slightly larger than dorsals; no pore-bearing, precloacal scales or precloacal depressions; forelimbs moderately long, slender; dorsal scales slightly raised, multicarinate; ventral scales of brachia smooth, raised, juxtaposed; scales beneath forearm smooth, raised, subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; sub digital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 12/46 196 Table 4 Mensural and meristic character states for the type series of Cnemaspis lineogularis sp. nov. All measurements taken are in millimeters and the abbreviations are defined in the materials and methods. BYU BYU ZMKU 62535 62536 R 00728 Holotype Paratype Paratype Supralabials 9 9 9 Infralabials 8 8 8 Ventral scales keeled (1) or smooth (0) 0 0 0 No. of precloacal pores 0 0 0 Precloacal pores continuous (1) or separated (0) / / / Precloacal pores elongate (1) or round (0) / / / No. of paravertebral tubercles 13 13 13 Tubercles linearly arranged (1) or more random (0) 1 1 1 Tubercles present (1) or absent (0) on lower flanks 0 0 0 Lateral caudal furrows present (1) or absent (0) 1 / 1 Caudal tubercles in lateral furrow (1) or not (0) 0 / 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 / 1 Lateral caudal tubercle row present (1) or absent (0) 1 / 1 Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) 0 / 1 Subcaudals keeled (1) or smooth (0) 0 / 0 Single median row of keeled subcaudals (1) or smooth (0) 0 / 0 Caudal tubercles encircle tail (1) or not (0) 0 / 0 Enlarged median subcaudal scale row (1) or not (0) 1 / 1 No. of postcloacal tubercles in males 1 1 / Enlarged femoral scales present (1) or absent (0) 1 1 1 Shield-like subtibial scales present (1) or absent (0) 0 0 0 Subtibial scales keeled (1) or smooth (0) 0 0 0 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 0 No. of 4th toe lamellae 29 27 29 SVL 38 35 29 TL 48 b 24 TW 2.9 / 2.66 FL 6.5 6.3 4.34 TBL 7.3 7.1 5.27 AG 17.5 15.7 10.6 HL 9.6 9.6 5.95 HW 6.4 6.3 5 HD 3.7 3.56 3.1 ED 2.5 1.78 1.71 EE 2.68 2.9 2.19 ES 5 4.2 3.54 EN 3.7 3.2 2.88 IO 3.1 2.5 1.99

(continued on next page)

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 13/46 197 Table 4 (continued) BYU BYU ZMKU 62535 62536 R 00728 Holotype Paratype Paratype EL 0.7 0.3 0.4 IN 1.2 1.2 0.89 Sex m m f

Notes. m, male; f, female; /, data unavailable or absent; b, broken. granular, lamellae of distal phalanges wide; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales raised, multicarinate, juxtaposed; dorsal scales on anterior margin of thighs enlarged, multicariante, becoming smaller posteriorly; ventral scales of thigh smooth; subtibial scales smooth, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; toes increase in length from first to fourth with fourth and fifth equal in length; 29,28 (R,L) subdigital lamellae on fourth toe; caudal scales similar to dorsal scale size, enlarge caudal tubercles arranged in segmented whorls, no encircling tail; caudal scales keeled, juxtaposed anteriorly; shallow, middorsal furrow; deeper, single, lateral furrow; enlarged, median, subcaudal scales; subcaudals smooth; median row of enlarged, keeled, subcaudal scales; transverse, tubercle rows do not encircle tail; caudal tubercles absent from lateral furrow; 2,1 (R,L) enlarged, postcloacal tubercles on lateral surface of hemipenal swellings at base of tail; posterior 30% of tail regenerated. Coloration. In life, dorsal ground color of head light beige-green, that of the body, limbs and tail slightly lighter than head; top of the head bearing, small black and light green markings; thin diffuse broken dark-brown to black postorbital stripe, extending to the nape; two dark lines radiating distally from orbit; dark paravertebral markings extend from nape to anterior fourth of tail where they transform into diffuse incomplete bands, intermixed with sage colored paravertebral blotches; single dark prescapular blotch dorsoanteriorly of forelimb insertion; limbs slightly lighter than dorsal ground color with randomly placed, diffuse dark blotches; all ventral surfaces grayish white, except gular region and anteriormost portion of throat orange with black midgular stripe and adjacent black stripes along the mandibular margin; posterior margin of orange gular coloration edged with black, transverse markings (Figs. 3–5). Variation. Paratypes approximate the holotype (BYU 62535) in general aspects of coloration except that the female paratype (ZMKU R 00728) lacks the black markings in the gular region and the yellowish-orange gular coloration is less prominent, additionally the dorsal coloration is much lighter. Selected body measurements and variation in squamation are presented in Table 4.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 14/46 198

) kamolnorranathi*

no / no no no no no no no no no no var no no no no no vandeventeri

no yes no no no no no no no no no no yes no no no no no punctatonuchalis yes yes no no no no no no yes no no no no no no no no no

(continued on next page roticanai

yes yes no no no no no no no no no no yes no no no no no omari

no / no no no no no no no no no no yes no no no no no

group chanardi no yes no no no no no no no no no no yes no no no no no

siamensis huaseesom yes yes yes no no no no no no no no no no yes yes no no no

Grismer et al., 2014d. siamensis

yes no no no no no no no no no no no no no no no no no

p nov. sp. thachanaensis

yes no no no no no no no no no no no no no no no no yes

p nov. sp. phangngaensis

yes no no no no no no no no no no yes? no no no no no yes from one another following tucdupensis

no no no no no no no no no no no no no no no no no no nuicamensis no no no no no no no no no no no no no no no no no no

Cnemaspis caudanivea no no no no no no no no no yes no no no no no no no no

group aurantiacopes

yes no no no no no no no no no no no no no no yes no yes laoensis*

/ no no no no no no no no no no no no no no no no / neangthyi

no no no no no no no no no no no no no no no no no no chanthaburiensis chanthaburiensis

yes no no no no no no no no no no no no no no no no no

p nov. sp. lineogularis yes no no no no no no no no no no no no no no no no yes Diagnostic color pattern characters separating various species of Thin yellow reticulum on side of neck Yellowish, prescapular crescent Forelimbs yellow Hind limbs yellow Reddish blotches or bands on limbs Forearms and forelegs orange Dorsal ground color magenta Ventral pattern sexually dimorphic Head yellow Reddish blotches on head and body Dense yellow reticulum on occiput and side of neck Ocelli on occiput and nape Ocelli on shoulder Dual ocelli on shoulder Ocelli on brachium and side of neck Thin, white, nuchal loop Large, black round spots on nape and anterior of body Dorsal color pattern sexually dimorphic Table 5

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 15/46

199

i* kamolnorranath

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o

vandeventeri s next

on

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o punctatonuchali

n o n o n o n o n o n o n o n o n o / / ye s n o n o n o n o

roticanai (continued
page)

n o n o n o n o n o n o n o n o n o ye s n o ye s n o n o n o n o omari

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o chanardi n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o

siamensis
group
huaseesom

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o va r n o

siamensis

n o n o n o ye s n o n o n o n o n o n o n o n o n o n o n o n o nov.

is sp. thachanaens

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o nov.

is sp. phangngaens

/ / n o ye s n o n o n o n o n o n o n o n o n o ye s n o n o tucdupensis

n o n o n o ye s ye s n o n o n o n o n o ye s n o n o n o n o n o

nuicamensis

n o n o n o ye s n o n o n o n o n o n o n o n o n o n o n o n o

caudanivea

n o ye s n o n o n o n o n o n o n o / / ye s n o n o n o n o

aurantiacopes

n o n o n o ye s n o n o n o n o n o n o n o n o n o n o n o ye s

laoensis*

n o n o n o n o n o n o n o n o n o / / ye s n o n o n o n o

neangthyi s

chanthaburiensis
group n o n o n o n o n o ye s n o n o n o n o n o n o n o n o n o n o chanthaburiensi

n o n o n o n o n o n o n o n o n o n o n o ye s n o n o n o n o nov.

s sp. lineogulari n o n o n o n o n o n o n o n o n o n o n o n o n o n o n o n o spots
spots tail bars
on
caudal
tail
tail
with
round
and
white
brown
tail
tail
bands
postscapular
or
dorsal
ground
light,
squarish,
yellow
flanks
black
vertebral
present Regenerated
yellow Regenerated
orange White,
caudal
tubercles Caudal
Wide
yellow
bands
Thin,
bands
anteriorly Uniform
ground
color Light
stripe Yellow
band Black,
paired,
paravertebral
dorsal
markings Small,
on
flanks Black
distinct
yellowish
Yellow
on
flanks Original
yellow Original
orange Dorsal
reddis color h (continued) Table
5

Wood
Jr
et
al.
(2017),
PeerJ,
DOI
10.7717/ 16/4 peerj.2884 6

200 kamolnorranathi*

no no no no no no no no no no no no no no no no no no no vandeventeri

no no no yes no no no no no var no yes yes no yes no no yes no punctatonuchalis

no no no / / / no yes / / / / / / / / no yes no roticanai

no no no no yes no no no yes no yes no no no no yes yes no no omari no no no no yes no no no no no yes no no no no yes yes no no

group chanardi no no no no yes no no no no no yes no no no no yes yes no no

siamensis huaseesom

no no no no yes no yes no yes no no no no yes no no yes no no siamensis

no no no no no no no no no no yes yes yes no yes no no no no

p nov. sp. thachanaensis

no no no no no no no no no yes no yes no yes no no no no no

p nov. sp. phangngaensis

no no no yes no no no / / no yes no no no no no yes no no tucdupensis

no yes no no no no no yes no yes no no no yes no yes no yes yes nuicamensis

no no no no yes no no no no no no yes no no no no no no no caudanivea no no no no no yes yes no no no no no no no no no no no no

group aurantiacopes

no yes no no no no no no no yes no yes no yes no yes no yes yes laoensis*

/ / / / / no no no no / / / / / / / / / / neangthyi

no no no no no no no no no no no no no no no no no no no chanthaburiensis chanthaburiensis

no no no no yes no no no no yes no no no yes no no no yes no

p nov. sp. lineogularis no no no no no no no no no yes no yes no no no no no no no ) (continued /, data unavailable; *, indicate species that are not included in the molecular analyses. Ventral surfaces of fore- limbs yellow Ventral surfaces of hind limbs orange Ventral surfaces of hind limbs yellow Subcaudal region yellow Subcaudal region orange At least posterior half of subcaudal region white Disticnt black and white caudal bands at least poste- riorly Gular region orange Gular region yellow Lineate gular markings Throat yellow Throat orange Pectoral region yellow Pectoral region orange Abdomen yellow Abdomen orange Ventral surfaces of fore- limbs orange Posterior portion of original tail white Posterior portion of origi- nal tail black Notes. Table 5

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 17/46

201 ) tucdupensis 51.0 8–10 7–9 0 0 / 16–22 w,1 1 1 0 1 0 0 0 0 0 w / (continued on next page 48.2 7–9 6–7 0 3–6 0,1 16–21 1 1 1 0, ant 1 0, ant 0 0 0 0 1 0 nuicamensis group. caudanivea 47.2 8,9 7,8 0 0–2 0,1 20–24 1 0 1 0, ant 1 0, ant 0 0 0 0 0,w 0 chanthaburiensis 58.4 9–11 8–10 0 0 / 23–31 1 1 1 0 1 1 0 0 0 0 1 / aurantiacopes in the 40.9 9 7 0 / / 22 0 1 1 1 0 0 0 0 0 0 w / laoensis* Cnemaspis neangthyi 0 20–26 1 1 1 1 1 1 0 0 0 0 1 54.0 11–13 10–12 0 2 1 from species of 21–25 0 42.2 8–10 7–10 0 6–9 chanthaburiensis 0,1 1 1 1 1 0 1 0 0 0 0, post, w 1 C. lineogularis lineogularis sp. nov. / 13 1 0 1 0 1 1 0 0 0 0 1 38 9 8 0 0 / Diagnostic morphological characters separating Caudal tubercles restricted to a single paravertebral row on each side (1)(0) or not Subcaudals keeled (1) or smooth (0) Single median row of keeled subcaudals (1) or smooth (0) Caudal tubercles encircle tail (1) or not (0) Enlarged median subcaudal scale row (1) or not (0) Precloacal pores elongate (1) or round (0) No. of paravertebral tubercles Tubercles linearly arranged (1) or more random (0) Tubercles present (1) or absent (0)lower on flanks Lateral caudal furrows present (1) or absent (0) Caudal tubercles in lateral furrow (1) or not (0) Ventrolateral caudal tubercles anteriorly (1) or not (0) Lateral caudal tubercle row present (1)absent or (0) Maximum SVL Supralabials Infralabials Ventral scales keeled (1) or smooth (0) No. of precloacal pores Precloacal pores continuous (1) or separated (0) Table 6

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 18/46

202 11 0–3 0 1 26–32 0 0 tucdupensis 2–4 0 0 27–33 10 0 0 nuicamensis 9 1,2 0,w 0,w 23–30 0 1 caudanivea 1,2 1 1 27–31 0 0 17 aurantiacopes 2,3 1 0 29 0 0 1 laoensis* 5 1 1 0 22–25 0 0 neangthyi 1–3 0,1 0 22–29 0 0 8 chanthaburiensis 3 1 0 0 27–29 1 0 lineogularis sp. nov. Grismer et al. (2014d). ) =) (continued w, weak; ant,Character anterior; abbreviations follow post, those of posterior; *, species that are not included in the molecular analyses; /, data unavailable or absent. No. of postcloacal tubercles in males Enlarged femoral scales present (1) or absent (0) Shield–like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) or smooth (0) Enlarged submetatarsal scales on 1st toe (1) or not (0) No. of 4th toe lamellae Sample size (n Table 6 Notes.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 19/46

203 Figure 6 (A) habitat and (B) microhabitat of Cnemaspis lineogularis sp. nov.

Etymology. The specific epithet lineogularis is derived from the Latin adjective linues for the word ‘‘line’’ and the nominative form of the Latin word gulare meaning ‘‘throat’’ and is in reference to the multiple dark gular lines present in the males of this species. Distribution. Only known from the type locality but we hypothesize it will be found in nearby karst formations (Figs. 1 and 6). Natural history. The type series and several other individuals were active during the day in shaded areas and would rapidly retreat to nearby cracks and crevices at the slightest provocation. We hypothesize this may be due to high predation as we found Trimeresurus fucatus in an ambush posture in the same microhabitat. No individuals were seen deep within the caves and from our observations, it appears this species primarily inhabits the more exterior surfaces of the karst tower (Fig. 6). The karst formations in this area are extensive and we assume this species has a much wider distribution than that reported here. We hypothesize that diurnality in this species is to avoid competition with and predation from the much larger Cyrtodactylus somroiyot with which it is hypothesized to be syntopic with. This is a commonly observed pattern among syntopic pairs of Cnemaspis

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 20/46 204 and Cyrtodactylus throughout their distributions in Southeast Asia (Grismer et al., 2014d, and references therein). Comparisons. Cnemaspis lineogularis sp. nov. can be differentiated for all other species in the chanthaburiensis group based on the following morphological and color pattern characteristics (see Tables 5 and 6 for additional comparisons). Cnemaspis lineogularis sp. nov. differs from C. chanataburiensis, C. neangthyi, C. laoensis, C. aurantiacopes, C. caudanivea, C. nuicamensis, and C. tucdupensis by having a smaller maximum SVL (38 mm vs. 42.2 mm, 54.0 mm, 40.9 mm, 58.4 mm, 47.2 mm, 48.2 mm, and 51.0 mm, respectively), by having less paravertebral tubercles (13 vs. 21–25, 20–26, 22, 23–31, 20–24, 16–21, and 16–22 respectively), and by having enlarged femoral scales. Cnemaspis lineogularis sp. nov. is further differentiated from C. neangthyi by having less supralabial scales (9 vs. 11–13). Cnemaspis lineogularis sp. nov. differs from C. neangthyi by having less infralabial scales (8 vs. 10–12) and from C. nuicamensis by having more infralabial scales (8 vs. 6–7). It is further differentiated from C. chanthaburiensis, C. neangthyi, C. aurantiacopes, C. caudanivea, and C. nuicamensis by lacking precloacal pores. From C. loaensis, C. lineogularis sp. nov. differs by having linearly arranged tubercles versus randomly arranged tubercles. Cnemaspis lineogularis sp. nov. differs from C. chanthaburiensis, C. neangthyi, C. laoensis, C. aurantiacopes, C. nuicamensis, and C. tucdupensis by lacking tubercles on the lower flanks. Cnemaspis lineogularis sp. nov. differs from C. chanthaburiensis, C. neangthyi, C. laoensis, by lacking caudal tubercles in the lateral furrow. Cnemaspis lineogularis sp. nov. has ventrolateral caudal tubercles anteriorly which separates it from C. chanthaburiensis and C. laoensis which lack this character. Cnemaspis lineogularis sp. nov. differs from C. loaensis, C. caudanivea, C. nuicamensis, and C. tucdupensis by the presence of a lateral caudal tubercle row. From C. chanthaburiensis, C. laoensis, C. caudanivea, and C. tucdupensis, C. lineogularis sp. nov. differs by having an enlarged median subcaudal scale row. C. lineogularis sp. nov. differs from C. laoensis and C. nuicamensies by having one postcloacal tubercle in males versus 2,3 and 2–4 respectively. C. lineogularis sp. nov. is further differentiated from C. caudanivea by lacking shield-like subtibial scales. Cnemaspis lineogularis sp. nov. differs from C. neangthyi, C. laoensis, and C. aurantiacopes by lacking keeled subtibial scales. Cnemaspis lineogularis sp. nov. differs from C. aurantiacopes and C. tucdupensis by lacking an enlarged submetatarsal scale on the 1st toe. Cnemaspis lineogularis sp. nov. is further differentiated from from C. neangthyi by having more 4th toe lamellae (27–29 vs. 22–25). Cnemaspis lineogularis sp. nov. is further differentiated from all other species in the chanthaburiensis group based on squamation and color pattern characteristics (Tables 5 and 6).

Cnemaspis phangngaensis sp. nov. urn:lsid:zoobank.org:act:6053C709-A409-4F65-B15C-8C647D7EDF1C Common name: The Phangnga Rock Gecko (Figs. 7–9)

Holotype. BYU 62538 adult male, collected at Phung Chang Cave, Mueang Phangnga District, Phangnga Province, Thailand (8.442344N, 98.514869E; 12 m a.s.l.), 26 July 2016, by PLW, LLG, CA, MC, MSG, MLM.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 21/46 205 Figure 7 (A) adult male holotype BYU 62538 and (B) female paratype BYU 62537 of Cnemaspis phangn- gaensis sp. nov.

Paratopotype. BYU 62537 adult female paratype bears all the same collection and locality information as the holotype. Diagnosis. Cnemaspis phangngaensis sp. nov. is distinguished from all other species of Cnemaspis in the siamensis group by the combination of the following morphological and color pattern characteristics: maximum SVL 42 mm; 10 supralabials; 10 infralabials;

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 22/46 206 Figure 8 Ventral coloration and sexual dichromatism of Cnemaspis phangngaensis sp. nov. (A) male holotype BYU 62538 and (B) female paratype BYU 62537.

ventral scales keeled; four continuous precloacal scales bearing a single round pore in males; 22 paravertebral tubercles linearly arranged; no tubercles on the lower flanks; lateral caudal furrows present; no caudal tubercles in the lateral furrows; lateral caudal tubercle row present; ventrolateral caudal tubercles anteriorly; caudal tubercles not encircling tail; caudal tubercles restricted to a single paravertebral row; subcaudals keeled bearing a single median row of enlarged keeled scales; two post cloacal tubercle in males; no enlarged femoral, tibial, or sub metatarsal scales; subtibials keeled; no enlarged median subcaudal scale row; no submetatarsal scale on first toe; 29 subdigital fourth toe lamellae; no enlarged median subcaudal scale row; dorsal and ventral color pattern sexually dimorphic; yellow or white bars present on flanks; prescapular marking present; anterior gular region dark yellowish, no dark lineate markings in males or females, and no mid-gular marking; posterior gular region and pectoral region whitish in males; abdomen yellow; subcaudal region yellow (Tables 5–7).

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 23/46 207 Figure 9 Dorsal coloration of the type series of Cnemaspis phangngaensis sp. nov. (A) male holotype BYU 62538 and (B) female paratype BYU 62537.

Description of the holotype. Adult male; SVL 42 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.27), somewhat narrow (HW/SVL 0.16), flattened (HD/HL 0.35), head distinct from neck; snout moderate (ES/HL 0.44), slightly concave in lateral view; postnasal region concave medially; scales on rostrum smooth becoming keeled posteriorly, raised, larger than conical scales on occiput; weak to absent supra ocular ridges; frontalrostralis sulcus shallow; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.20); extra-brillar, fringe scales largest anteriorly; pupil round; ear opening more oval, taller than wide; rostral slightly concave, dorsal 80% divided by longitudinal median groove; rostral bordered posteriorly by supra nasals and one small azygous scale and laterally by first supralabials; 10, 10 (R,L) slightly raised supralabials decreasing in size posteriorly; 10, 10 (R,L) infralabials decreasing in size posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered by small postnasal scales; mental large, triangular,

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 24/46 208 concave, bordered posteriorly by three postmentals; gular and throat scales raised, keeled, small and round. Body slender, elongate (AG/SVL 0.45); small, raised, keeled, dorsal scales equal in size throughout body, intermixed with several large , multicarinate conical tubercles more or less randomly arranged; tubercles extend from the occiput to base of the tail; no tubercles on flanks; pectoral and abdominal scales keeled, not larger posteriorly; abdominal scales slightly larger than dorsals; two pore-bearing, continuous, precloacal pores on each side; forelimbs moderately long, slender; dorsal scales slightly raised, keeled; ventral scales of brachia smooth, raised, juxtaposed; scales beneath forearm smooth, slightly raised, subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; sub digital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales raised, multicarinate, juxtaposed; ventral scales of thigh, slightly raised, conical, keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; no enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; toes increase in length from first to fourth with fourth and fifth equal in length; 29, 29 (R,L) subdigital lamellae on fourth toe; caudal scales similar to dorsal scale size, enlarge caudal tubercles arranged in segmented whorls, not encircling tail; caudal scales keeled, juxtaposed anteriorly; shallow, middorsal furrow; deeper, single, lateral furrow; enlarged, median, subcaudal scales; subcaudals keeled; median row of enlarged, keeled, subcaudal scales; transverse, tubercle rows do not encircle tail; caudal tubercles absent from lateral furrow; 1, 1 (R,L) enlarged flat, postcloacal tubercle on lateral surface of hemipenal swellings at base of tail; posterior 30% of tail missing. ⇠ Coloration. In life dorsal ground color of head light beige, that of the body, limbs and tail slightly darker than the head with darker irregular blotches; top of the head bearing, small black and sage markings; thin diffuse broken dark brown to black postorbital stripe, extending to the nape, not complete; light sage vertebral blotches extending form the nape to tail where they transform into diffuse near complete irregular bands; intermixed with light sage blotches; single light-yellowish prescapular crescent dorsoanteriorly of forelimb insertion; flanks with irregular incomplete sage to yellowish-orange bars becoming more orange distally; limbs slightly darker than dorsal ground color with randomly placed, diffuse dark and sage colored blotches; all ventral surfaces grayish-white, except gular, abdominal, and subcaudal regions are yellowish-orange, with more pronounced darker yellow stippling (Figs. 7–9). Variation in the type series. The female paratype (BYU 62537) approximates the holotype in general aspects of coloration except the overall dorsal coloration is lighter and the ventral

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 25/46 209 Table 7 Mensural and meristic character states for the type series of Cnemaspis phangngaensis sp. nov. All measurements are taken in millimeters and the abbreviations are defined in the materials and methods. BYU BYU 62538 62537 holotype paratype Supralabials 10 10 Infralabials 10 10 Ventral scales keeled (1) or smooth (0) 1 1 No. of precloacal pores 4 0 Precloacal pores continuous (1) or separated (0) 1 / Precloacal pores elongate (1) or round (0) 0 / No. of paravertebral tubercles 22 22 Tubercles linearly arranged (1) or more random (0) 1 1 Tubercles present (1) or absent (0) on lower flanks 0 0 Lateral caudal furrows present (1) or absent (0) 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 1 Caudal tubercles restricted to a single paravertebral row on 11 each side (1) or not (0) Subcaudals keeled (1) or smooth (0) 1 1 Single median row of keeled subcaudals (1) or smooth (0) 1 1 Caudal tubercles encircle tail (1) or not (0) 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 0 No. of postcloacal tubercles in males 2 / Enlarged femoral scales present (1) or absent (0) 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 Subtibial scales keeled (1) or smooth (0) 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 0 0 No. of 4th toe lamellae 29 30 SVL 42 41 TL 23b 44 TW 3.3 3.2 FL 6.37 6.6 TBL 8.26 8.23 AG 19.28 17.6 HL 11.6 11.1 HW 6.79 6.56 HD 4.1 4.1 ED 2.4 2.4 EE 3.1 3.1 ES 5.17 4.86 EN 3.9 4.2 IO 2.6 2.9

(continued on next page)

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 26/46 210 Table 7 (continued) BYU BYU 62538 62537 holotype paratype EL 1 0.99 IN 2.85 2.75 Sex m f

Notes. m, male; f, female; /, data unavailable or absent; b, broken. coloration is a uniform light yellow and is not as prominent in the gular and abdominal regions. Select body measurements and variation in squamation are presented in Table 7. Etymology. The specific epithet phangngaensis is a noun in apposition to the type locality where this species is found. Distribution. Only known from the karst formation in which it is found, the Phung Chang Cave, Phangnga, Mueang Phangnga, Thailand. We hypothesize that this species will be found on nearby contiguous karst formations. Natural history. Cnemaspis phangngaensis inhabits a karst formation in a lowland limestone forest (Fig. 10) surrounded by highly disturbed, urbanized habitat. The male holotype was collected at night on the karst approximately 15 m above the ground on the exterior surface of the tower and the female was collected at night sleeping on a leaf approximately 1.2 m above the limestone forest floor adjacent to the nearby karst formation. Individuals were also observed active during the day, but avoided being caprtured be retreating into the rock crevices. We hypothesize that these are diurnal karst dwellers that use the vegetation at night for refuge. We hypothesize that diurnality in this species is to avoid competition with and predation from the much larger Cyrtodactylus lekaguli with which it is syntopic. Comparisons. The phylogenetic analysis recovers the chanardi group and C. phangngaensis sp. nov. as the sister species to a clade containing C. omari and C. roticani (Fig. 2). This relationship is further supported by the following derived morphological characters (sensu Grismer et al., 2014d), prescapular crescent present, yellow abdomen, yellow ventral surfaces of the hind limbs and tail being yellow and numerous other morphological and color pattern characteristics (Tables 5 and 8). C. phangngaensis sp. nov. differs from C. chanardi, C. omari, and C. roticanai by having; more infralabial scales (10 vs. 6–8, 7,8, and 7,8, respectively); continuous precloacal pores; paravertebral tubercles linearly arranged; lacking tubercles on the lower flank; ventrolateral caudal tubercles anteriorly; caudal tubercles restricted to a single paraveterbral row on each side; a single median row of keeled subcaudals. Cnemaspis phangngaensis sp. nov. is further differentiated from C. chanardi and C. omari by having a larger maximum SVL (42 mm vs. 40.1 mm and 41.3 mm, respectively). Cnemaspis phangngaensis sp. nov. differs from C. omari, and roticani by having more supralabial scales (10 vs. 8,9 and 8,9, respectively). C. phangngaensis sp. nov. differs from C. chanardi by having fewer precloacal pores (4 vs. 6–8). Cnemaspis phangngaensis sp. nov. differs from C. roticani by having fewer paravertebral tubercles (22 vs. 25–27). From C. roticanai, C. phangngaensis sp. nov. differs by lacking caudal tubercles in the lateral furrow and by having a lateral caudal tubercle row present. Cnemaspis phangngaensis sp. nov. differs from C. omari by lacking caudal tubercles encircling the tail

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 27/46 211 ) kamolnorranathi* 0 1 w 0 w 37.8 8,9 7,8 0,w 7 1 1 19–24 w 1 1 1 0 1 (continued on next page 0 1 w 0 1 1 0 0 0 0 1 0 0 1 vandeventeri 44.7 8,9 7–9 4 25–29 punctatonuchalis 0 0 0 0 1 49.6 8 7,8 0 0 / / 24–27 w 1 1 0 1 ant group. 0 1 0 0 w 1 0 0 0 1 1 1 0 0 roticanai 47.0 8,9 7,8 3–6 25–27 siamensis omari 0 1 0 1 0 41.3 8,9 7,8 1 4 0 0 22–29 w,0 w,1 1 0 0 1 0 1 0 0 1 1 0 0 20–30 0 1 1 0 0 1 chanardi 40.1 7–10 6–8 6–8 from one another in the huaseesom 0 0 0 0 0 43.5 7–10 6–9 0 5–8 1 0 18–24 w,0 1 1 1 0 0 Cnemaspis 0 1 0 0 1 39.7 8,9 6–8 1 0 / / 19–25 0 1 1 0 0 1 siamensis 1 1 1 0 0 1 / / 1 1 1 0 1 1 thachanaensis sp. nov. 39 10,11 9–11 0 15–19 1 1 1 0 0 0 22 1 0 1 0 1 1 42 10 10 1 4 1 phangngaensis sp. nov. Diagnostic morphological characters separating species of Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) Subcaudals keeled (1) or smooth (0) Single median row of keeled subcaudals (1) or smooth (0) Caudal tubercles encircle tail (1) or not (0) Enlarged median subcaudal scale row (1) or not (0) Precloacal pores elongate (1) or round (0) No. of paravertebral tubercles Tubercles linearly arranged (1) or more random (0) Tubercles present (1) or absent (0) on lower flanks Lateral caudal furrows present (1) or absent (0) Caudal tubercles in lateral furrow (1) or not (0) Ventrolateral caudal tubercles anteriorly (1) or not (0) Lateral caudal tubercle row present (1) or absent (0) Maximum SVL Supralabials Infralabials Ventral scales keeled (1) or smooth (0) No. of precloacal pores Precloacal pores continuous (1) or separated (0) Table 8

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 28/46

212 24–28 1,2 0 0,1 0 3 kamolnorranathi* 0 1–3 0 1 0 vandeventeri 0 24–28 3 29–31 1–3 0 0 0 5 punctatonuchalis 0 1,2 0 1 0 roticanai 0 26–29 8 25–28 4 1 0 1 0 omari 0 25–30 1 0 1 0 chanardi 0 25 1,2 0 0 0 0 21–31 5 huaseesom 24–26 1,2 1 0 12 0 siamensis 0 0 1 1 0 0 24 6 thachanaensis sp. nov. Grismer et al. (2014d). 29 2 1 0 2 0 phangngaensis sp. nov. 0 ) (continued w, weak; ant,Character anterior; abbreviations follow post, that of posterior; *, species that are not included in the molecular analyses; /, data unavailable or absent. No. of postcloacal tubercles in males Enlarged femoral scales present (1) or absent (0) Shield–like subtibial scales present (1) or absent (0) Subtibial scales keeled (1) or smooth (0) Enlarged submetatarsal scales on 1st toe (1) or not (0) No. of 4th toe lamellae Sample size Notes. Table 8

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 29/46

213 Figure 10 (A) general karst and limestone forest near the type locality of Cnemaspis phangngaensis sp. nov. (B) karst microhabitat where C. phangngaensis occurs.

and by having more lamellae under the 4th toe (29 vs. 25–28). Cnemaspis phangngaensis sp. nov. is further differentiated from C. chanardi by lacking an enlarged median subcaudal scale row. From C. chanardi and C. omari, C. phangngaensis differs by have two postcloacal tubercles in males versus one. Cnemaspis phangngaensis is further differentiated from all other species in the siamensis group based on squamation and color pattern characteristics (Tables 5 and 8).

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 30/46 214 Cnemaspis thachanaensis sp. nov. urn:lsid:zoobank.org:act:3581C94E-6170-4F42-9159-E2B564B576F1 Common name: The Tha Chana Rock Gecko (Figs. 11–13) Cnemaspis kamolnorranathi (Grismer et al., 2010, pg. 29) Cnemaspis kamolnorranathi (Grismer et al., 2014d, pg. 130)

Holotype. BYU 62544 adult male, collected at Tham Khao Sonk hill, Tha Chana District, Surat Thani Province, Thailand (9.549878N, 99.175544E; 107 m a.s.l.), 30 July 2016, by PLW, LLG, CA, MC, MSG, MLM. Paratopotypes. All paratypes (BYU 62542–62543, ZMKU R 00729–00731) bear the same collection and locality data as the holotype. Diagnosis. Cnemaspis thachanaensis sp. nov. is distinguished from all other species of Cnemaspis in the siamensis group by the combination of the following morphological and color pattern characteristics: maximum SVL 39 mm; 10 or 11 supralabials; 9–11 infralabials; ventral scales keeled; no precloacal pores in males; 15–19 paravertebral tubercles linearly arranged; tubercles generally present on the lower flanks; lateral caudal furrows present; no caudal tubercles in the lateral furrows; ventrolateral caudal tubercles anteriorly; presence of lateral caudal tubercle row; caudal tubercles not encircling tail; caudal tubercles restricted to a single paravertebral row; subcaudals keeled bearing a single median row of enlarged keeled scales; one or two post cloacal tubercles in males; no enlarged femoral or tibial scales; subtibials keeled; enlarged submetatarsal scale on first toe; 23–25 subdigital fourth toe lamellae; sexually dimorphic for ventral and dorsal coloration; yellow or white bars present on flanks; prescapular marking present; gular region yellowish-orange, dark incomplete lineate markings in males, less prominent in females; abdomen, limbs and subcaudal region whitish (Table 9). Description of the holotype. Adult male; SVL 33 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.29), somewhat narrow (HW/SVL 0.16), flattened (HD/HL 0.37), head distinct from neck; snout moderate (ES/HL 0.44), snout slightly concave in lateral view; postnasal region concave medially; scales on rostrum smooth becoming keeled posteriorly, raised, larger than conical scales on occiput; weak to absent supra ocular ridges; frontalrostralis sulcus shallow; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.22); extra-brillar, fringe scales largest anteriorly; pupil round; ear opening more oval than round, taller than wide; rostral slightly concave, dorsal 80% divided by longitudinal median groove; rostral bordered posteriorly by supra nasals and one small azygous scale and laterally by first supralabials; 11, 11 (R,L) slightly raised supralabials decreasing in size posteriorly; 10, 10 (R,L) infralabials decreasing in size posteriorly; nostrils elliptical, oriented dorsoposteriorly; bordered by small postnasal scales; mental large, triangular, concave, bordered posteriorly by three postmentals; gular scales small, smooth, raised and round; throat scales subimbricate, keeled, small and round. Body slender, elongate (AG/SVL 0.44); small, raised, keeled, dorsal scales equal in size throughout body, intermixed with several large, multicarinate conical tubercles more or less randomly arranged; tubercles extend from the occiput to base of the tail; enlargerd

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 31/46 215 Figure 11 Coloration of Cnemaspis thachanaensis sp. nov. (A) male holotype BYU 62544 and (B) BYU 62542 female paratype.

multicarinate conical tubercles on flanks; pectoral and abdominal scales keeled, not larger posteriorly; abdominal scales slightly larger than dorsals; no pore-bearing, precloacal pores on either side; forelimbs moderately long, slender; dorsal scales slightly raised, keeled; ventral scales of brachia smooth, raised, juxtaposed; scales beneath forearm smooth, slightly raised, subimbricate; palmar scales smooth, juxtaposed, raised; digits long with an inflected joint; claws recurved; sub digital lamellae unnotched; lamellae beneath first phalanges granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; fingers increase in length from first to fourth with fourth and fifth equal in length; hind limbs slightly longer and thicker than forelimbs; dorsal scales raised, multicarinate, juxtaposed; ventral scales of thigh, slightly raised, conical, keeled; subtibial scales keeled, flat, imbricate, with no enlarged anterior row; plantar scales smooth, juxtaposed, raised; enlarged submetatarsal scales beneath first metatarsal; digits elongate with an inflected jointed; claws recurved; subdigital lamellae unnotched; lamellae beneath first phalanges

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 32/46 216 Figure 12 Ventral coloration and sexual dichromatism of the type series of Cnemaspis thachanaensis sp. nov., males: (A) BYU 62543, (B) BYU 62544 (holotype), (C) ZMKU R 00731, females: (D) ZMKU R 00729, (E) ZMKU R 00730, (F) BYU 62542.

Figure 13 Dorsal coloration of the type series of Cnemaspis thachanaensis sp. nov., males: (A) BYU 62543, (B) BYU 62544 (holotype), (C) ZMKU R 00731, females: (D) ZMKU R 00729, (E) ZMKU R 00730, (F) BYU 62542.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 33/46 217 Table 9 Menusural and meristic character state for the type series of Cnemaspis thachanaensis sp. nov. All measurements are taken in millime- ters and the abbreviations are defined in the materials and the methods. BYU ZMKU R BYU BYU ZMKU R ZMKU R 62544 00731 62543 62542 00729 00730 holotype paratype paratype paratype paratype paratype Supralabials 10 11 10 10 10 10 Infralabials 10 11 10 10 9 9 Ventral scales keeled (1) or smooth (0) 1 1 1 1 1 1 No. of precloacal pores 0 0 0 / / / Precloacal pores continuous (1) or separated (0) / / / / / / Precloacal pores elongate (1) or round (0) / / / / / / No. of paravertebral tubercles 15 19 15 17 15 16 Tubercles linearly arranged (1) or more random (0) 1 1 1 1 1 1 Tubercles present (1) or absent (0) on lower flanks 1 1 0 1 1 1 Lateral caudal furrows present (1) or absent (0) 1 1 1 1 1 1 Caudal tubercles in lateral furrow (1) or not (0) 0 0 0 0 0 0 Ventrolateral caudal tubercles anteriorly (1) or not (0) 1 1 1 1 1 1 Lateral caudal tubercle row present (1) or absent (0) 1 1 1 1 1 1 Caudal tubercles restricted to a single paravertebral row on 111111 each side (1) or not (0) Subcaudals keeled (1) or smooth (0) 1 1 1 1 1 1 Single median row of keeled subcaudals (1) or smooth (0) 1 1 1 1 1 1 Caudal tubercles encircle tail (1) or not (0) 0 0 0 0 0 0 Enlarged median subcaudal scale row (1) or not (0) 0 0 0 0 0 0 No. of postcloacal tubercles in males 0 0 0 / / / Enlarged femoral scales present (1) or absent (0) 1 1 1 0 0 0 Shield-like subtibial scales present (1) or absent (0) 0 0 0 0 0 0 Subtibial scales keeled (1) or smooth (0) 1 1 1 1 1 1 Enlarged submetatarsal scales on 1st toe (1) or not (0) 1 1 1 1 1 1 No. of 4th toe lamellae 24 23 24 25 23 23 SVL 33 37 34 39 35 35 TL 41 44r 43 46 6b 16b TW 3.6 3.91 3.32 4 3.4 3.7 FL 5.97 5.68 5.39 5.83 5 5.9 TBL 6.8 7.3 6.19 7.14 6.9 6.5 AG 14.55 16 13.1 17.72 14.49 15.6 HL 9.79 10.3 9.74 11.49 9.37 9.8 HW 5.55 6.27 5.3 6.6 5.45 5.7 HD 3.64 4.1 3.8 4.32 3.9 3.8 ED 2.21 2.4 2 2.36 2.04 1.9 EE 2.85 2.79 2.6 2.79 2.67 3 ES 4.4 4 3.6 4.29 4.29 4.57 EN 2.5 1.18 2.6 3.1 2.79 3.7 IO 2.58 2.97 2.3 3.29 3.1 2.8 (continued on next page)

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 34/46 218 Table 9 (continued) BYU ZMKU R BYU BYU ZMKU R ZMKU R 62544 00731 62543 62542 00729 00730 holotype paratype paratype paratype paratype paratype EL 0.76 0.82 0.67 1 0.71 0.75 IN 2.1 2.8 2.4 2.2 2.29 2.2 Sex m m m f f f Notes. m, Male; f, Female; /, Data unavailable or absent; b, Broken; r, regenerated. granular proximally, widened distally; lamellae beneath phalanx immediately following inflection granular, lamellae of distal phalanges wide; interdigital webbing absent; toes increase in length from first to fourth with fourth and fifth equal in length; 24,24 (R,L) subdigital lamellae on fourth toe; caudal scales similar to dorsal scale size, enlarge caudal tubercles arranged in segmented whorls, not encircling tail; caudal scales keeled, juxtaposed anteriorly; shallow, middorsal furrow; deeper, single, lateral furrow; enlarged, median, subcaudal scales; subcaudals keeled; median row of enlarged, keeled, subcaudal scales on last 2/3 of tail; transverse, tubercle rows do not encircle tail; caudal tubercles absent from lateral furrow; 1,1 (R,L) enlarged flat, postcloacal tubercle on lateral surface of hemipenal swellings at base of tail. Coloration. In life dorsal ground color of head light-brown, that of the body, limbs and tail slightly darker than the head with even darker irregular blotches; top of the head bearing, small dark-brown and light-green markings; thin diffuse broken dark brown to black postorbital stripe, extending to the nape, not complete; light-green vertebral blotches extending form the nape to tail where they transform into diffuse near complete irregular bands intermixed with dark brown blotches turning into bands posteriorly; flanks with irregular incomplete small light-green colored blotches to yellow-orange bars becoming smaller posterior; limbs much lighter than dorsal ground color, limbs grayish-white and dark brown incomplete irregular bands; all ventral surfaces grayish-white, except gular and throat regions are yellow-orange not restricted to the gular region and extend onto the throat and the anterior region of the pectoral region in males, incomplete transverse markings in the gular region in male and is less prominent in females (Figs. 11–13). Variation. The paratypes approximate the holotype (BYU 62544) in general aspects of morphology except that the female paratypes lack precloacal pores and yellow-orange gular regions. Paratypes ZMKU R 00731, BYU 62542, and BYU 62541 have more paravertebral tubercles (19, 17, 16 respectively vs. 15), dark irregular gular spots not as prominent in females (Fig. 12). Select body measurements and additional variation in squamation are presented in Table 9. Etymology. The specific epithet thachanaensis is a noun in apposition to the type locality where this species is found. Distribution. This species is only know from the type locality Thom Sonk Hill, Tha Chana District, Surat Thani Province, Thailand and we expect that it will be found on nearby adjacent karst formations (Fig. 14).

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 35/46 219 Figure 14 (A) karst and limestone forest near the type locality of Cnemaspis thachanaensis sp. nov. (B) karst microhabitat of Cnemaspis thachanaensis sp. nov.

Natural history. Cnemaspis thachanaensis inhabits a karst tower embedded within a highly disturbed lowland limestone forest. One male individual was observed during the day situated upside down on a karst overhang displaying its yellow-orange throat by doing push-ups. All other specimens were found active during the day on the karst and we hypothesize that these are diurnal karst dwellers. No specimens were observed at night. Grismer et al. (2010) noted that one specimen (CUMZ-R 2009,624-3) was collected on a vine near the adjacent limestone. Karst dwelling species of Cnemaspis have been know to sleep on vegetation at night (Grismer et al., 2010; Grismer et al., 2014d, P Wood, pers. obs., 2016). This species may use the vegetation at night for refuge to avoid Cyrtodactylus thirakaputhi which is nocturnal and maybe a potential predator. Remarks. Specimen CUMZ-R 2009,6,24-3 was collected from Thom Sonk Hill, Tha Chana District, Surat Thani Province and was described as C. kamolnorranathi in Grismer et al. (2010). Grismer et al. (2010) noted that the relatively wide separation ( 110 km) ⇠ between the type locality of C. kamolnorranathi (Petchphanomwat Waterfall, Tai Rom Yen National Park, Ban Nasan District, Surat Thai Province) and the locality of the paratype

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 36/46 220 CUMZ-R 2009,6,24-3 from Thom Sonk Hill, Tha Chana District, Surat Thani Province and suggested that there are probably undiscovered, geographically intervening populations in the appropriate habitat separating these two localities (Grismer et al., 2014d). Grismer et al. (2010) and Grismer et al. (2014d) noted that there is exceptional intrapopulational variation in the degree of keeling of the ventral and the subtibial scales in C. kamolnorranathi suggesting the possibility that C. kamolnorranathi may be composed of multiple species. After examining additional specimens from Thom Sonk Hill, Tha Chana District, Surat Thani Province (BYU 62542, ZMKU R 00729–00731 and the paratype CUMZ-R 2009,6,24- 3) we determined that CUMZ-R 2009,6,24-3 is not conspecific with C. kamolnorranathi and with additional specimens it can be diagnosed as a new species (see comparisons below for details). Here we remove CUMZ-R 2009,6,24-3 from C. kamolnorranthi and place it in C. thachanaensis sp. nov. restricting C. kamolnorranathi to the Petchphanomwat Waterfall, Tai Rom Yen National Park, Ban Nasan District, Surat Thai Province. There are no genetic samples of C. kamolnorranthi available to further test this hypotheses, however we present strong morphological evidence separating these species. Comparisons. Cnemaspis thachanaensis sp. nov. is the sister species to a clade containing C. siamensis and C. vandeventeri (Fig. 2). Although we were not able to obtain genetic material for C. kamolnorranathi we compare it here using morphology to demonstrate that the paratype (CUMZ-R 2009,6,24-3, MS101) is conspecific with C. thachanaensis sp. nov. Cnemaspis thachanaensis sp. nov. differs from C. siamensis and C. vandeventeri by having a smaller SVL (39 mm, vs. 39.7 mm and 44.7 mm) and by having a larger maximum SVL from C. kamolnorranathi (39 mm vs. 37.8 mm). C. thachanaensis sp. nov. differs from C. siamensis, C. vandeventeri, and C. kamolnorranathi by; having more supralabial scales (10–11 vs. 8–9, 8–9, 8–9, respectively); having more infralabials (9–11 vs. 6–8, 7–9, and 7–8, respectively); having paravertebral tubercles linearly arranged; having ventrolateral caudal tubercles anteriorly; having caudal tubercles restricted to a single paravertebral row on each side; having a single median row of keeled subcaudal scales; lacking a single enlarged subcaudal scale row; lacking postcloaclal tubercles in males; the presence of an enlarged submetatarsal scale on the 1st toe. Cnemaspis thachanaensis sp. nov. is further differentiated from C. kamolnorranathi by having keeled ventral scales. Cnemaspis thachanaensis sp. nov. differs from C. vandeventeri and C. kamolnorranathi by lacking precloacal pores. We can further differentiate C. thachanaensis sp. nov. from C. vandeventeri by having less paravertebral tubercles (15–19 vs. 25–29). Cnemaspis thachanaensis sp. nov. differs from C. kamolnorranathi by lacking tubercles in the lateral furrow. Cnemaspis thachanaensis sp. nov. is further differentiated the more distantly related species C. huaseesom and C. punctatonuchalis in the siamensis group by having a smaller maximum SVL (39 mm vs. 43.5 mm and 49.6 mm, respectively); having more supralabials 10,11 vs. 8; having caudal tubercles restricted to a single paravertebral row; having keeled ventral scales; single median row of keeled subcaudals; lacking enlarged median subcaudal scale row; by lacking postcloacal tubercles in males. Cnemaspis thachanaensis sp. nov. differs by having more infralabials 9–11 vs. 7, 8 in C. punctatonuchalis. Cnemaspis thachanaensis sp. nov. differs from C. huaseesom by lacking precloacal pores. From C. huaseesom, C. thachanaensis sp. nov. differs by having ventrolateral caudal tubercles anteriorly and the presence of a lateral

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 37/46 221 caudal tubercle row. Cnemaspis thachanaensis sp. nov.differs from C. punctatonuchalis by having keeled subcaudal scales. Cnemaspis thachanaensis sp. nov. differs from C. huaseesom by having keeled subtibial scales an an enlarged submetatarsal scale on the first toe. From C. punctatonuchalis, C. thachanaensis sp. nov. differs by having less fourth toe lamellae, 24 vs. 29–31. Cnemaspis thachanaensis sp. nov. is differentiated from all other species in the siamensis group based on squamation and color pattern characteristics (Tables 5 and 8).

DISCUSSION The discovery of three new species of karst-dwelling Cnemaspis from Peninsular Thailand is not surprising, given the nature of the vastly unexplored karst and limestone forests dispersed throughout this area. Peninsular Malaysia received considerable attention with respect to herpetofaunal surveys, yet new karst-dwelling species are being discovered and described every year (see Grismer et al., 2016a, for a summary). The results of these surveys have resulted in the discovery of 14 species of geckos (including Cnemaspis and Cyrtodactylus as well as two snakes Grismer et al. (2016a)). In comparison, Peninsular Thailand has received little attention with most of the focus on the genus Cyrtodactylus resulting in the discovery and description of 15 species in the last 55 years, with 14 of these being described in the last 15 years (see Table 6 in Grismer et al., 2016a). However, there has been limited field research on the Thai karst-dwelling Cnemaspis from these areas (Grismer et al., 2010). With the small amount of time spent in Phangnga, Tha Chana, and Prachuap Khiri Khan, we were able to discover three new species (C. lineogularis sp. nov., C. phangngaensis sp. nov., C. thachanaensis sp. nov.) and successfully collect genetic samples of C. punctatonuchalis and C. vandeventeri. We expect that as more time is focused collecting specimens from the unexplored karst formations additional new species will be discovered. From the fieldwork that has been conducted on the Thai and Malaysian karst formations a fair amount of Cyrtodactylus and Cnemaspis have been discovered and described. On some of these formations both the nocturnal Cyrtodactylus and diurnal Cnemaspis occur syntopically. For example both Cyrtodactylus lekaguli and Cnemaspis phangngaensis sp. nov. at Phung Chang Cave, Phangnga, Thailand, both Cyrtodactylus astrum and Cnemaspis omari in Perlis, Malaysia, and both Cyrtodactylus langkawiensis and Cnemaspis roticani on Pulau langkawi, Malaysia. Comparing the phylogenetic relationships of these Cyrtodactylus with the Cnemaspis reveals an identical phylogeographic pattern (Phangnga (Perlis, Pulau Langkawi)). Corroborating these relationships are the calculated average mean pairwise sequence divergence within the respective genera (Cyrtodactylus 9.4% (Grismer et al., 2016b) and Cnemaspis 9.6% [this study]). From these preliminary analyses we hypothesize that the formation of these karst formations may have been the resulting factor for simultaneous speciation events within these two respective genera. With additional fieldwork and data collection in these areas more detailed analyses with divergence times can be estimated to investigate the timing of these divergence events. The inclusion of C. punctatonuchalis and C. vandeventeri in the phylogenetic analyses helps test previous morphological hypotheses set forth by Grismer et al. (2010) and

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 38/46 222 Grismer et al. (2014d), and has also contributed towards a more complete phylogeny of the genus Cnemaspis (49 of the 55 named species including the three new species described herein). Cnemaspis punctatonuchalis was nested within the siamensis group confirming the placement solely based on morphological and color pattern characteristics by Grismer et al. (2014d), which was also hypothesized to be more closely related to the other northern species (north of the Isthmus of Kra, C. huaseesom and C. siamensis). This is further supported here as the sister species to C. huaseesom (Fig. 2). Cnemaspis vandeventeri was hypothesized based on its distribution that it should align with the siamensis group, however Grismer et al. (2014d) also suggested that the presence of a light prescapular crescent that diagnoses a monophyletic group composed of C. chanardi, C. phangngaensis, C. omari and C. roticanai may suggest that it is more closely related to this group. The phylogenetic placement of C. vandeventeri is well nested in the siamensis group confirming the placement based on its distribution of Grismer et al. (2010) and Grismer et al. (2014d), however the hypothesis that it may be more closely related to the group with the prescapular crescent is not supported by our phylogenetic hypothesis and could represent an instance of convergent evolution of the prescapular crescent. This in not surprising based on the well documented parallel/convergent evolution present in the genus Cnemaspis (Grismer et al., 2014d), and further analyses to address hypotheses pertaining to parallel/convergent evolution of multiple traits are in preparation (P Wood et al., 2016, unpublished data). The phylogenetic position of C. lineogularis as the sister taxon to the entire chanthaburiensis group, indicates a trans-Gulf of Thailand relationship with other species from southern Indochina. This is not a novel biogeographic pattern and the close relationship between Indochinese and Malaysian lineages has been observed in Butterfly lizards in the genus Leiolepis (Grismer et al., 2014a), and in some species of Cyrtodactylus (Grismer et al., 2015). However, the previous documented cases of this pattern are much further south on the peninsula. This pattern could easily be explained by previous cyclic sea level fluctuations that exposed the Sunda Shelf providing multiple dispersal corridors between the Thai-Malay Peninsula and Indochina (e.g., Voris, 2000; Sathiamurthy & Voris, 2006; Woodruff, 2010). Further investigation into the biogeographic patterns for Cnemaspis are in preparation (P Wood et al., 2016, unpublished data), and with the continued discovery of new species of Cnemasapis in the area, these broader studies will contribute to the understanding of the complex biogeography patterns on the Thai-Malay Peninsula. The discovery of three new species of Cnemaspis described here underscores the need for additional fieldwork in the karst towers of the Thai-Malay Peninsula and the surrounding areas to aid in conservation efforts, document the herpetofauna diversity, and provide data for biogeographic studies.

ACKNOWLEDGEMENTS We are thankful for general discussions and logistical conversations with Attapol Rujirawan (aka Bank), Siriporn Yodthong, Natee Ampai, Korkhwan Termprayoon, Piyawan Phuanprapai, and Sengvilay Seateun. We would also like to thank Todd R. Jackman, Angelica Crottini, and D. James Harris for providing feedback that greatly improved the quality of the manuscript.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 39/46 223 ADDITIONAL INFORMATION AND DECLARATIONS

Funding Funding was from the Department of Biology at Brigham Young University. Additional funding for this research is from the NSF dimensions grant EF-1241885 issued to JWS and the Doctoral Dissertation Improvement Grant (DDIG #1501198) issued to PLWJ and JWS. Partnerships for Enhanced Engagement in Research (PEER) Science program (grant PGA-2000003545), which is a partnership between the US Agency for International Development (USAID) and the National Science Foundation (NSF), provided funding for AA. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Grant Disclosures The following grant information was disclosed by the authors: Department of Biology at Brigham Young University. NSF dimensions: EF-1241885. Doctoral Dissertation Improvement: DDIG #1501198. Enhanced Engagement in Research (PEER) Science program: PGA-2000003545. US Agency for International Development (USAID). National Science Foundation (NSF).

Competing Interests The authors declare there are no competing interests.

Author Contributions Perry Lee Wood Jr conceived and designed the experiments, performed the experiments, • analyzed the data, wrote the paper, prepared figures and/or tables, reviewed drafts of the paper. L. Lee Grismer conceived and designed the experiments, performed the experiments, • wrote the paper, reviewed drafts of the paper. Anchalee Aowphol, César A. Aguilar, Micheal Cota, Marta S. Grismer and Matthew L. • Murdoch performed the experiments, reviewed drafts of the paper. Jack W. Sites Jr contributed reagents/materials/analysis tools, reviewed drafts of the • paper.

Animal Ethics The following information was supplied relating to ethical approvals (i.e., approving body and any reference numbers): Brigham Young University’s Institutional Animal Care and Use Committee (IACUC) has approved the animal use protocol for this study (protocol # 160401).

Field Study Permissions The following information was supplied relating to field study approvals (i.e., approving body and any reference numbers):

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 40/46 224 We received a collecting permit from the National Research Council Thailand (NRCT) to PLW and AA (No. 0002/7606). DNA Deposition The following information was supplied regarding the deposition of DNA sequences: GenBank accession numbers KY091231–KY091244. New Species Registration The following information was supplied regarding the registration of a newly described species: Publication ID: urn:lsid:zoobank.org:author:E2AF9554-5062-48C6-84AA-FA50D720DEF7 Family: Gekkonidae, Genus: Cnemaspis, Species: lineogularis urn:lsid:zoobank.org:act:8E3B21A4-93BF-4D08-B8D1-0A3EEF6BE44F Family: Gekkonidae, Genus: Cnemaspis, Species: phangngaensis urn:lsid:zoobank.org:act:6053C709-A409-4F65-B15C-8C647D7EDF1C Family: Gekkonidae, Genus: Cnemaspis, Species: thachanaensis urn:lsid:zoobank.org:act:3581C94E-6170-4F42-9159-E2B564B576F1 Supplemental Information Supplemental information for this article can be found online at http://dx.doi.org/10.7717/ peerj.2884#supplemental-information.

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Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 42/46 226 morphological and molecular analyses with descriptions of seven new species. Zootaxa 3520:1–55. Grismer LL, Wood Jr PL, Shahrul A, Awal R, Norhayati A, Muin M, Sumontha M, Grismer J, Chan K, Quah ES, Pauwels O. 2014d. Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa 3880(1):1–147 DOI 10.11646/zootaxa.3880.1.1. Grismer LL, Wood Jr PL, Shahrul A, Grismer MS, Quah ESH, Murdoch ML, Muin MA, Davis HR, Aguilar C, Klabacka R, Cobos AJ, Aowphol A. 2016b. Two new Bent- toed Geckos of the Cyrtodactylus pulchellus complex from Peninsular Malaysia and multiple instances of convergent adaptation to limestone forest ecosystems. Zootaxa 4105(5):401–429 DOI 10.11646/zootaxa.4105.5.1. Grismer LL, Wood Jr PL, Shahrul A, Quah ES, Muin MA, Mohamed M, Onn CK. 2014e. The phylogenetic relationships of three new species of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae) from poorly explored regions in northeastern peninsular Malaysia. Zootaxa 3786(3):359–381 DOI 10.11646/zootaxa.3786.3.6. Grismer LL, Wood Jr PL, Tri N, Murdoch ML. 2015. The systematics and independent evolution of cave ecomorphology in distantly related clades of Bent-toed Geckos (Genus Cyrtodactylus Gray, 1827) from the Mekong Delta and islands in the Gulf of Thailand. Zootaxa 3980(1):106–126 DOI 10.11646/zootaxa.3980.1.6. Holloway JD. 1986. Origins of lepidopteran faunas in high mountains of the Indo- Australian tropics. In: High altitude tropical biogeography. Oxford: Oxford University Press, 533–566. Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP. 2001. Bayesian inference of phylogeny and its impact on evolutionary biology. Science 294(5550):2310–2314 DOI 10.1126/science.1065889. Hughes JB, Round PD, Woodruff DS. 2003. The Indochinese–Sundaic faunal transition at the Isthmus of Kra: an analysis of resident forest bird species distributions. Journal of Biogeography 30(4):569–580 DOI 10.1046/j.1365-2699.2003.00847.x. Jenkins PD, Kilpatrick CW, Robinson MF, Timmins RJ. 2005. Morphological and molecular investigations of a new family, genus and species of rodent (Mammalia: Rodentia: Hystricognatha) from Lao PDR. Systematics and Biodiversity 2(4):419–454 DOI 10.1017/S1477200004001549. Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T, Ashton B, Meintjes P, Drummond A. 2012. Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28(12):1647–1649 DOI 10.1093/bioinformatics/bts199. Kiew R. 1991. The limestone flora. In: The State of Nature Conservation in Malaysia. Kuala Lumpur: Malaysian Nature Society, 42–50. Kiew R. 1998. Limestone, quartzite and ultramafic vegetation. In: The Encyclopedia of Malaysia: plants. Singapore: Editions Didier Miller, 26–27.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 43/46 227 Komo I. 1998a. The karst morphology of Langkawi. In: The Encyclopedia of Malaysia: the environment. Singapore: Editions Didier Miller, 40–41. Komo I. 1998b. Caves and cave systems: the Mulu caves. In: The Encyclopedia of Malaysia: the environment. Singapore: Editions Didier Miller, 42–43. Latinne A, Waengsothorn S, Herbreteau V, Michaux JR. 2011. Evidence of complex phylogeographic structure for the threatened rodent Leopoldamys neilli, in Southeast Asia. Conservation Genetics 12(6):1495–1511 DOI 10.1007/s10592-011-0248-3. Macey JR, Larson A, Ananjeva NB, Fang Z, Papenfuss TJ. 1997. Two novel gene orders and the role of light-strand replication in rearrangement of the verte- brate mitochondrial genome. Molecular Biology and Evolution 14(1):91–104 DOI 10.1093/oxfordjournals.molbev.a025706. Maddison W, Maddison D. 2015. Mesquite: a modular system for evolutionary analysis. version 3.04. 2015. Available at http://mesquiteproject.org. Ng P. 1991. Cancrocaeca xenomorpha, new genus and species, a blind troglobitic fresh- water hymenosomatid (Crustacea: Decapoda: Brachyura) from Sulawesi, Indonesia. Raffles Bulletin of Zoology 39(1):59–73. Parnell J. 2013. The biogeography of the Isthmus of Kra region: a review. Nordic Journal of Botany 31(1):001–015 DOI 10.1111/j.1756-1051.2012.00121.x. Patou M-L, Chen J, Cosson L, Andersen D, Cruaud C, Couloux A, Randi E, Zhang S, Veron G. 2009. Low genetic diversity in the masked palm civet Paguma larvata (Viverridae). Journal of Zoology 278(3):218–230 DOI 10.1111/j.1469-7998.2009.00570.x. Pauwels O, David P, Chimsunchart C, Thirakhupt K. 2003. Reptiles of Phetchaburi Province, Western Thailand: a list of species, with natural history notes, and a discussion on the biogeography at the Isthmus of Kra. The Natural History Journal of Chulalongkorn University 3(1):23–53. Pauwels OS, Sumontha M, Latinne A, Grismer LL. 2013. Cyrtodactylus sanook (Squa- mata: Gekkonidae), a new cave-dwelling gecko from Chumphon Province, southern Thailand. Zootaxa 3635(3):275–285 DOI 10.11646/zootaxa.3635.3.7. Posada D, Crandall KA. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14(9):817–818 DOI 10.1093/bioinformatics/14.9.817. Raes N, Cannon CH, Hijmans RJ, Piessens T, Saw LG, Van Welzen PC, Slik JF. 2014. Historical distribution of Sundaland’s Dipterocarp rainforests at Quaternary glacial maxima. Proceedings of the National Academy of Sciences of the United States of America 111(47):16790–16795 DOI 10.1073/pnas.1403053111. Ronquist F, Teslenko M, Van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP. 2012. Mrbayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3):539–542 DOI 10.1093/sysbio/sys029. Sacha M. 2015. Herpetoreisen in Thailand: Erste Eindrücke aus Krabi. Sauria 37(3):43–55.

Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 44/46 228 Sathiamurthy E, Voris HK. 2006. Maps of Holocene sea level transgression and sub- merged lakes on the Sunda Shelf. The Natural History Journal of Chulalongkorn University 2(Supplement):1–43. Schilthuizen M. 2004. Land snail conservation in Borneo: limestone outcrops act as arks. Journal of Conchology Special Publication 3:149–154. Schilthuizen M, Liew T-S, Elahan BB, Lackman-Ancrenaz I. 2005. Effects of karst forest degradation on pulmonate and prosobranch land snail communities in Sabah, Malaysian Borneo. Conservation Biology 19(3):949–954 DOI 10.1111/j.1523-1739.2005.00209.x. Schilthuizen M, Vermeulen J, Davison G, Gittenberger E. 1999. Population structure in a snail species from isolated Malaysian limestone hills, inferred from ribosomal DNA sequences. Malacologia 41:271–284. Siler CD, Oaks JR, Esselstyn JA, Diesmos AC, Brown RM. 2010. Phylogeny and bio- geography of Philippine Bent-toed Geckos (Gekkonidae: Cyrtodactylus) contradict a prevailing model of Pleistocene diversification. Molecular Phylogenetics and Evolution 55(2):699–710 DOI 10.1016/j.ympev.2010.01.027. Stamatakis A. 2006. Raxml-vi-hpc: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22(21):2688–2690 DOI 10.1093/bioinformatics/btl446. Stamatakis A, Hoover P, Rougemont J. 2008. A rapid bootstrap algorithm for the raxml web servers. Systematic Biology 57(5):758–771 DOI 10.1080/10635150802429642. Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. 2013. MEGA6: molecu- lar evolutionary genetics analysis version 6.0. Molecular Biology and Evolution 30(12):2725–2729 DOI 10.1093/molbev/mst197. Tija HD. 1998. Limestone and Karst morphology. In: The Encyclopedia of Malaysia: the environment. Singapore: Editions Didier Miller, 38–39. Uetz P, Freed P, Ho≤ek J. 2016. The reptile database. Available at http://www.reptile- database.org/ (accessed on September 2016). Vermeulen J, Whitten T. 1999. Biodiversity and cultural property in the management of limestone resources. Washington, D.C.: World Bank, 120. Voris HK. 2000. Maps of pleistocene sea levels in southeast asia: shorelines, river systems and time durations. Journal of Biogeography 27(5):1153–1167 DOI 10.1046/j.1365-2699.2000.00489.x. Wilcox TP, Zwickl DJ, Heath TA, Hillis DM. 2002. Phylogenetic relationships of the dwarf boas and a comparison of bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution 25(2):361–371 DOI 10.1016/S1055-7903(02)00244-0. Wood Jr PL, Heinicke MP, Jackman TR, Bauer AM. 2012. Phylogeny of Bent-toed Geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution 65(3):992–1003 DOI 10.1016/j.ympev.2012.08.025. Wood Jr PL, Quah ES, Anuar S, Muin MA. 2013. A new species of lowland karst dwelling Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from northwestern Peninsular Malaysia. Zootaxa 3691(5):538–558 DOI 10.11646/zootaxa.3691.5.2.

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Wood Jr et al. (2017), PeerJ, DOI 10.7717/peerj.2884 46/46 230 Chapter 5: A Geologically-driven Rapid Radiation on the Sunda Shelf

231 A Geologically-driven Rapid Radiation on the Sunda Shelf

1, 2 3 4 5
Perry
L.
Wood
Jr ⇤,
Adam
Leache´ ,
Tony
Gamble ,
Anchalee
Aowphol ,
L.
Lee
Grismer &
Jack

W.
Sites
Jr1

1Department
of
Biology
and
Bean
Life
Science
Museum,
Brigham
Young
University,
Provo,
Utah

84602
USA.

2Department
of
Biology
and
Burke
Museum
of
Natural
History
and
Culture,
University
of
Wash- ington,
Seattle,
WA,
98195
USA.

3Department
of
Biological
Sciences,
Marquette
University,
Milwaukee,
WI
53201,
USA.

4Kasetsart
University,
Faculty
of
Science,
Department
of
Zoology,
Chatuchak,
Bangkok,
10900,

Thailand.

5Department
of
Biology,
La
Sierra
University,
4500
Riverwalk
Parkway,
Riverside,
CA,
92515

USA.

"CTUSBDU

The
geological
complexity
and
the
understudied
biodiversity
hot
spot
of
Southeast
Asia
pro- vides
outstanding
options
for
studying
patterns
and
processes
of
diversification
and
evolution.
Here
we
provide
the
most
complete
phylogeny
of
rock
geckos
based
on
a
genomic
dataset,
coupled
with
well
informed
paleogeographic
reconstructions,
to
generate
hypotheses
about
the
tempo
and
modes
of
diversification
in
this
species-rich
clade
on
the
Sunda
Shelf.

We
demonstrate
that
rock
geckos
had
a
significant
rate
shift
in
speciation
correlating
to
dynamic
geological
episodes
on
the
Sunda
Shelf.
We
show
that
the
temporal
radiation
of
rock
geckos
coincides
with
the
temporal
diversification
of
other
terrestrial
vertebrates
across

Sundaland.

232 *OUSPEVDUJPO

Southeast
Asia
(SEA)
is
a
complex,
intriguing
and
vastly
understudied
region
of
the
world1.

The
continental
and
insular
landmasses
of
SEA
comprise
4%
of
the
earth’s
landmasses,
yet
con- tain
20–25%
of
the
earth’s
biodiversity,
making
the
Sundaic
region
(Sundaland)
one
of
the
world’s
biodiversity
hotspots2–10.
Sundaland
is
comprised
of
the
Sunda
Shelf
and
the
Thai-Malay
Penin- sula,
the
large
islands
of
Borneo,
Java,
Sumatra,
and
many
smaller
archipelagos
(Figure
S1).
One
of
the
most
prominent
features
of
Sundaland
is
the
Sunda
Shelf,
a
vast
area
of
continental
shelf
composed
of
the
Sunda
Plains
(currently
submerged
beneath
the
South
China
Sea),
and
the
sur- rounding
land-positive
fringe
regions
of
the
Thai-Malay
Peninsula,
Sumatra,
Borneo,
and
Java.

Cyclical
climatic
and
sea
level
fluctuations
over
the
last
2.4
million
years
have
facilitated
range
expansion
and
contraction/isolation
of
biotas11,12,
especially
in
the
cooler
montane
rainforests13,14,
resulting
in
the
formation
of
interglacial
upland
refugia15,16.
Sea
level
fluctuations
have
also
driven
the
formation
and
isolation
of
island
archipelagos8,17,18.
Due
to
this
long
and
varied
geological
history,
the
majority
of
the
biota
of
this
vast
landscape
exists
in
a
refugial
state2,4,7,12,19.

The
older
geological
history
of
SEA
has
been
impacted
and
shaped
by
dynamic
Cenozoic
tectonic
forces20,21,
but
the
combined
influence
of
Cenozoic
geological
processes
and
Quaternary
climatic
changes
as
drivers
of
the
evolutionary
history
of
the
SEA
biota
is
less
clear
and
is
often
over-looked19,22,23.
Recent
advances
in
Cenozoic
paleogeographic
reconstructions
of
the
Sunda

Shelf20,21,24
provide
an
excellent
framework
to
test
a
priori
hypotheses
about
the
temporal/spatial
drivers
of
diversification
throughout
this
region.

Geckos of the genus Cnemaspis comprise a species-rich clade (55 species25) that is widely

233 distributed
throughout
Sundaland
(Figure
S1),
and
study
of
this
group
may
provide
general
in- sights
into
some
of
the
diversification
patterns
throughout
this
region.
Recent
molecular
phyloge- netic
studies
based
on
a
small
number
of
mitochondrial
and
nuclear
genes
recover
a
well-supported
monophyletic
genus
and
resolve
“recent
history”
relationships
within
some
groups,
but
they
lack
sufficient
signal
to
resolve
the
deeper
nodes
in
this
clade25–27.
The
distribution
of
Cnemaspis
throughout
Sundaland,
its
well-supported
monophyly,
low
vagility,
and
unresolved
deep
relation- ships,
make
this
clade
a
model
system
to
test
multiple
nested
biogeographic
hypotheses.

Here
we
use
a
genomic
dataset
to
construct
a
time-calibrated
Cnemaspis
phylogeny,
and
ad- dress
the
following
questions:
(1)
are
the
unresolved
polytomies
in
the
genus
“hard”
or
“soft”?
(2)
is
there
evidence
for
gradual
or
punctuated
increases
in
the
accumulation
of
species
in
Cnemaspis?

And
(3)
do
dynamic
geological
events
across
the
Sunda
Shelf
at
different
time
scales
coincide
with
patterns
of
lineage
diversification
in
Cnemaspis?

Results

The
maximum
likelihood
(ML)
analyses
for
the
concatenated
datasets
(514,
440,
and
105
loci)
composed
of
the
50p,
60p,
and
75p
all
recover
Cnemaspis
as
monophyletic.
Nodal
support
varied
between
the
different
ML
analyses
(Figure
S2).
For
example,
the
50p
dataset
had
a
longer
align- ment
(267,912
bp/514
loci)
compared
to
the
60p
(231,027
bp/440
loci),
and
75p
(57,940bp/105
loci),
but
had
missing
individuals
per
locus
compared
to
the
other
two
datasets.
The
50p
dataset
recovered
a
topology
with
higher
support
values
than
the
60p
and
75p
datasets
(Figure
S2),
which

234 is
congruent
with
the
results
of
other
studies28,
so
we
present
these
results
here,
which
generally
recovers
a
topology
concordant
with
similar
levels
of
nodal
support
as
the
60p
and
75p
datasets
and
analyses.
A
few
well-supported
nodes
recovered
in
the
BI
analyses
are
not
supported
by
the

ML
analyses,
and
other
nodes
well-supported
by
ML
are
not
strongly
supported
in
our
BI
trees

(Figure
S3A).

The
concatenated
and
species
tree
analyses
from
A-II
and
SVDQ
recover
similar
topolo- gies
with
respect
to
some
other
well-supported
deeper
nodes,
although
some
weakly
supported
relationships
were
recovered
in
the
species
tree
analyses
(Figure
S3A,B,
S4).
Further,
some
inter- relationships
within
the
more
nested
(younger)
clades
are
incongruent
between
the
two
analyses.

Three
samples
(C.
affinis,
C.
hangus,
and
C.
narathiawatensis)
were
only
represented
by
a
single
gene
and
their
phylogenetic
placement
was
ambiguous
in
the
SVDQ
analysis,
therefore
they
were
removed,
but
were
included
in
the
A-II
analysis
which
recovered
the
same
phylogenetic
placement
of
these
three
species
as
the
previous
analyses
(Figure
S5).
The
quartets
generated
from
the
A-II
analyses
showed
that
68.42%
of
all
the
quartets
in
the
gene
trees
were
also
recovered
in
the
species
tree.

We
recovered
the
Ca
Mau
clade
as
sister
to
all
other
Cnemaspis
groups,
which
is
concor- dant
with
previous
studies25,26,
but
the
remaining
relationships
differ
from
previous
studies25,26.

We
recover
the
Southern
Sunda
clade
as
the
sister
group
to
the
remaining
species
of
Cnemaspis,
whereas
previous
studies
recovered
the
Pattani
clade
as
the
sister
group
to
all
other
Cnemaspis
groups
excluding
the
Ca
Mau
clade
(Figure
S3).
However
these
deeper
internodes
in
the
pre-

235 vious
studies
were
characterized
by
short
branch
lengths
with
no
significant
support.
Here
we
resolved
the
deeper
phylogenetic
relationships
in
the
concatenated
analyses
with
well-supported
nodes,
whereas
our
other
analyses
recovered
the
same
topology
albeit
with
lower
support
values
for
the
deeper
nodes
(i.e.,
PP=0.64,
BS=70,
and
SVDB=70
for
the
deepest
node
and
PP=0.92,
BS=80,
and
SVDB=51
for
the
other,
Figures
S4A,B).
To
further
investigate
this
issue
we
estimated
the
posterior
probabilities
at
these
problematic
nodes
and
calculated
the
percentages
of
the
quartets
in
the
gene
trees
that
agree
with
the
respective
branches
(“quartet
support”)
on
these
short
internal
nodes
in
the
backbone
of
the
Cnemaspis
tree
(Figure
S4).

To
further
investigate
these
two
problematic
nodes
we
estimated
the
posterior
at
these
prob- lematic
nodes
and
calculated
the
percentages
of
the
quartets
in
the
gene
trees
that
agree
with
the
respective
branches
(“quartet
support”)
on
these
short
internal
nodes
in
the
backbone
of
the
Cne- maspis
tree
(Figure
S4).

Our
divergence
analyses
with
the
fossil
and
the
root
height
calibrations
estimated
that
crown

Gekkota
diverged
from
its
common
ancestor
approximately
105.9
MYA,
which
is
well
within
the
estimates
of
previous
analyses29,30.
The
common
ancestor
of
the
Ca
Mau
clade
and
the
ancestor
to
all
other
Cnemaspis
species
diverged
during
the
mid-Paleocene
approximately
61.1
MYA
(Figure

1A,
S5).
Within
this
“all
other”
clade,
the
Southern
Sunda
clade
diverged
from
the
ancestor
of
the remaining
species
41.9
MYA
(Figure
1A,
S5).
The
former
ancestor
of
the
“remaining
species”
⇠ clade
then
diverged
into
the
Malaysian
Peninsula
clade,
while
the
ancestor
to
the
Pattani
and
the

Thai
Peninsula/Indochina
clade
diverged
approximately
39.2
MYA,
and
the
ancestor
to
the
(Pattani

236 +
Thai
Peninsula/Indochina)
clade
diverged
37.3
MYA.
In
summary,
all
major
clades
(Southern

Sunda(Malay
Peninsula(Pattani
+
Thai
Peninsula/Indochina))),
diverged
between
33.4–41.9
MYA

(41.9/39.2/37.3/33.4
MYA,
respectively).

BAMM31
diversification
analyses
show
a
relatively
high
early
average
speciation
rate
()
for
all
Cnemaspis,
increasing
from
2
to
5.9
species/my
(Figure
S6A);
exclusion
of
the
Ca
Mau
clade
increases
the
average
speciation
rate
slighty
from
5.9
to
6.37
species/my
(Figure
S6B).
There
was
a
positive
extinction
rate
of
µ=0.41
species/my
for
all
Cnemaspis,
and
a
slight
increase
in
extinction
around
60–40
MYA
(Figure
S6C).
Exclusion
of
the
Ca
Mau
clade
increases
this
extinction
rate
almost
imperceptibly
from
0.41
to
0.42,
so
extinctions
remain
relatively
constant
through
time

(Figure
S6D).
We
found
strong
evidence
for
a
single
rate
shift
in
speciation
along
the
branch
leading
to
the
species-rich
Cnemaspis
clade
that
is
the
sister
group
to
the
Ca
Mau
clade,
with
the
core
shift
probability
of
0.70
(Figure
1,
S7).
We
then
calculated
the
Bayes
Factor
(BF)
by
incorporating
the
rate
shift
at
the
branch
subtending
this
large
clade
under
the
prior
alone,
and
found
strong
evidence
(BF
=
40.9,
based
on
Kass
and
Raftery
199532)
for
a
rapid
radiation
“Out
of
the
Sunda
Shelf”
(OSS,
Figure
1).

To
further
investigate
the
OSS
idea,
we
estimated
a
LTT
plot
to
test
for
a
rapid
radiation
in

Cnemaspis,
and
estimated
net
diversification
rates
in
Geiger.
The
LTT
plot
showed
a
significant
departure
from
the
null
CR
model
of
speciation,
with
the
greatest
accumulation
of
species
during the
past
5–20
MYA
and
the
with
the
OSS
radiation
starting
much
earlier
(Figure
S8A).
This
result
⇠ has
a
significantly
negative -statistic
(=
-1.847611;
p
=
0.04)
based
on
the
20,000
simulated

237 pure-birth
constant
rate
trees
for
the
MCCR
test
(Figure
S8B);
this
is
evidence
of
a
significant
increase
in
diversification
rates
(rapid
radiation).

The
BioGeoBears
ancestral
biogeographical
range
reconstruction
analysis
selected
the
DI-

VALIKE+J
model
as
the
best
fit
to
our
data
(Table
S2),
and
recovers
an
ancestral
node
for
all

Cnemaspis
composed
of
(Ca
Mau
+
Borneo
+
Peninsular
Malaysia);
the
proto-Sunda
Shelf.
This
analysis
also
recovered
two
vicariant
and
twelve
jump-dispersal
founder
events.
The
ancestral
node
of
all
Cnemaspis
represents
a
vicariant
event
that
separated
the
Ca
Mau
area
and
Peninsu- lar
Malaysia+Borneo
61.1
MYA
(Figure
1).
Directly
following
the
separation
of
the
proto-Sunda

Shelf,
another
vicariant
event
split
the
Borneo
and
Peninsular
Malaysia
highland
areas
(41.9
MYA),
giving
rise
to
the
ancestral
range
for
the
Southern
Sunda
clade
and
the
ancestor
to
the
Peninsular

Malaysia
area.
Within
the
Southern
Sunda
clade
there
are
five
jump-dispersal
or
founder
events,
the
details
of
which
will
be
the
focus
of
another
paper
and
not
discussed
here.
The
Peninsular

Malaysia
area
has
a
founder
event/jump
dispersal
event
to
Central
Indochina
39.2
MYA,
giving
rise
to
the
Borneo
clade
(Figure
1).

Discussion.
Previous
phylogenetic
hypotheses
of
Cnemaspis25,26
had
unresolved
short
internodes,
which
obscured
the
relationships
of
the
larger
clades
and
limited
biogeographic
inferences
pertain- ing
to
temporal
patterns
of
diversification
across
the
Sunda
Shelf.
Here
we
resolve
the
deeper
nodes
of
the
phylogeny
with
moderate
to
strong
support
for
the
for
the
species
tree
analyses
(the
concate- nated
dataset,
Figure
S3).
Previous
studies25,26
recovered
four
major
clades:
Ca
Mau,
Pattani,

Southern
Sunda,
and
Northern
Sunda,
and
in
both
the
species
tree
and
the
concatenated
analyses,

238 we
recovered
the
Ca
Mau,
Pattani,
and
the
Southern
Sunda
clades,
but
also
a
paraphyletic
Northern

Sunda
group
sensu
Grismer
et
al.26
(Figure
S3A,B).
Here
we
designate
the
paraphyletic
North- ern
Sunda
group
into
two
new
clades,
the
Malaysian
Peninsula
and
the
Thai
Peninsula/Indochina
clades,
respectively
(Figure
S3A,B).

The
resolution
of
these
short
internal
branches
provide
evidence
that
this
is
not
a
“hard”
polytomy,
but
a
“soft”
polytomy
that
is
evident
of
a
rapid
diversification
within
Cnemaspis
“OSS”.

However,
the
concatenated
analyses
can
be
misleading
because
they
ignore
the
individual
gene
histories
and
can
even
provide
strong
support
for
the
wrong
tree33.
In
comparison
our
species
tree
analyses
recovered
identical
deeper
relationships
of
Cnemaspis
with
moderate
support
for
short
internal
branches.
The
quartet
approach
for
those
two
branches
indicate
that
a
small
percentage
of
the
genetree
quartets
are
congruent
with
the
branches
in
the
species
tree
(Figure
S3A,B);
this
is
ev- idence
of
high
levels
of
incomplete
lineage
sorting
(ILS),
which
is
expected
during
times
of
rapid
lineage
accumulation34,35.
ILS
topologies
(evidence
of
ancient
lineage
accumulations
associated
with
short
internodes)
have
been
documented
in
a
number
of
clades,
including
birds36,
rodents37,
and
cichlid
fishes38.
Our
LTT
plot
shows
a
statistically
significant
increase
in
speciation
rate
in
the
large
Cnemaspis
clade,
consistent
with
the
hypothesis
of
an
accelerated
lineage
accumulation from
40
MYA
throughout
the
clades
geologically
more
recent
history.
In
addition,
the
strong
⇠ Bayes
Factor
(40.9)
from
the
BAMM
analysis
and
70%
probability
(Figure
1A
and
S7)
recovered a
significant
rate
shift
along
the
lineage
leading
to
the
large
clade
of
Cnemaspis,
which
is
contem- poraneous
with
the
initial
increase
in
rapid
lineage
accumulation
from
the
MCCRT
and
visually
in
the
LTT
(Figure
S8A,B).

239 The
biogeographic
analyses
suggest
that
Cnemaspis
most
likely
originated
on
the
proto-

Sunda
Shelf
in
the
Paleocene
(Figure
1A)
and
rapidly
radiated
during
times
of
significant
Eocene
geological/vicariant
events
of
Eastern
Indochina
(Ca
Mau
clade)1,5,21,39.
These
analyses
also
sug- gest
that
there
were
two
major
vicariant
events
during
the
early
history
of
Cnemaspis;
one
each
during
the
Paleocene
and
the
late
Eocene
encompassing
the
vicariant
events
of
the
ancestral
proto-

Sunda
Shelf,
followed
by
the
separation
of
the
paleogeographic
Peninsula
Malaysian
and
Bornean
highlands5,21
(Figure
1C).
All
major
clades
of
Cnemaspis
except
for
the
Ca
Mau
clade,
then
di- versified
rapidly
between
41.9–37
MYA
during
the
separation
of
the
Malaysian
and
the
Bornean
highlands.
The
temporal
diversification
of
Cnemapsis
is
not
a
novel
event
and
many
other
flora
and
fauna
including
the
banana
family
(Musaceae)40,
megophryid
and
rhacophorid
frogs41,42,
dragon
lizards43,
geckos44,47,
and
the
evolution
of
volant
vertebrates
in
Southeast
Asia45,
have
revealed
similar
initial
diversification
conclusions
and
are
often
attributed
to
the
collision
of
the
Indian
plate
with
Eurasia.

The
initial
diversification
of
Cnemaspis
starts
during
the
Paleocene,
with
a
rapid
diversifi- cation
during
the
Eocene
and
the
LTT
plots
identifies
that
there
is
significant
increases
in
lineage
accumulation
during
the
more
recent
history
(Miocene-Eocene).
The
Oligocene–Miocene
diver- sification
( 20–30
My)
pattern
shown
in
the
LTT
plots
of
Cnemaspis
has
also
been
documented
⇠ in
a
number
of
other
squamate
reptiles,
including
agamid
(Wood
et
al.
in
prep),
gekkonids46,47, and
homolopsid
snakes48.
Other
groups,
including
some
plants
in
the
custard
apple
family
and
palms49,50,
spiny
frogs
42,51,
ranid
frogs52
and
marine
gastropods53,
birds54
also
experienced
in- creased
diversification
rates
in
SEA
during
the
Oligocene-Miocene
transition.
Some
hypotheses

240 for
these
shared
common
patterns
are
contentious
owing
to
competing
hypotheses
about
the
extent
of
connectivity
of
the
Sunda
Shelf
islands5,
or
its
division
by
marine
transgressions10.
However,
recent
evidence
favors
the
marine
transgressions10,55,56,
though
the
extent,
location
and
frequency
of
these
are
still
unclear48.
The
possibility
that
Cnemapsis
has
undergone
multiple
rapid
radia- tions
is
a
realistic
possibility
with
the
BAMM
analysis
recovering
an
older
rate
shift
in
speciation
and
the
LTT
plots
with
a
rapid
increase
in
speciation
during
the
Oligocene-Miocene
transition
and
provides
a
competing
hypothesis
for
the
the
rapid
radiation
of
Cnemaspis.

This
study
provides
a
well-resolved
phylogeny
for
nearly
all
species
of
Sundaland
Cnemaspis
and
presents
the
phylogeny
in
a
temporal
perspective
coupled
with
a
rapid
radiation
corresponding
to
major
geological
events
from
the
Paleocene
to
the
Miocene.
Cnemaspis
most
likely
originated
in
the
Paleocene
and
had
initial
rate
shifts
of
rapid
lineage
accumulation
starting
in
the
Eocene
with
additional
radiations
in
the
more
recent
history
during
the
Oligocene-Miocene
transition.
The
number
of
dynamic
geological
events
in
the
Paleocene
and
Miocene
across
the
Sunda
Shelf
region
provided
dispersal
corridors
at
different
time
intervals
facilitating
the
dispersal
and
rapid
speciation
of
Cnemaspis.
Further
generating
alternative
hypotheses
pertaining
to
the
patterns
and
processes
of
speciation
and
evolution
across
the
Sunda
Shelf.

Methods

Data
collection.
Complete
sampling
of
all
of
the
species
of
Cnemaspis
is
difficult
based
on
their
large
distribution
and
many
of
the
species
are
micro
endemics
with
no
available
genetic
material.

We
sampled
one
individual
per
species
for
almost
all
putative
species
of
Cnemaspis,
including

241 two undescribed species (n=53), but not including C. dringi, C. laoensis, C. sundagekko, and C. kamolnorranathi, for which genetic samples are unavailable. We included 21 outgroup taxa based on57 and a list of voucher specimens used in this study is presented in (Table S1).

Targeted sequence capture and UCE data collection. Targeted sequence capture data were col- lected using a dataset of RNA probes specific for iguanian lizards58. The commercially synthesized custom probes were designed to target 585 loci with 2X tiling (two 120 bp probes per locus) using

MYbaits target enrichment kit (MYcroarray Inc., Ann Arbor, MI, USA). These probes targeted 541 ultraconserved elements (UCEs) that were used in the Tetrapods-UCE-5Kv1 probes59 (describes in detail at http://www.ultraconserved.org), for details about the the probe design and selection see60.

In addition to the UCEs we added 44 loci from the Squamate Tree of Life project (SToL)61.

Genomic DNA was extracted from liver or skeletal muscle tissue stored in 95% ethanol

R using
the
animal
DNA
extraction
protocol
from
a
Qiagen
DNeasy
tissue
kit
(Valencia,
CA,
USA).
To
check
the
quality
of
the
DNA
we
ran
electrophoreses
gels
to
ensure
that
the
DNA
had a
high
molecular
weight.
All
DNA
samples
( 400
ng)
that
had
high
molecular
weights
were
⇠ then
sonicated
with
a
targeted
size
peak
of
400
bp
on
a
Bioruptor
Pico
(Diagenode
Inc.)
for
seven cycles
of
sonication
for
seven
seconds
then
ninety
seconds
of
rest.
We
then
prepared
DNA
libraries
using
the
Illumina
TruSeq
Nano
DNA
library
preparation
kit.
The
DNA
library
samples
were
then
hybridized
to
the
RNA-probes
in
presence
of
a
blocking
mixture
composed
of
forward
and
reverse
compliments
of
the
Illumina
TruSeq
Nano
adapters,
with
insoines
in
place
of
the
indices,
as
well
as
salmon
blocking
and
Chicken
blocking
mixtures
(Chicken
Hybloc,
Applied
Genetics
Lab
Inc.)
to
reduce
repetitive
DNA
binding
to
beads.
Libraries
were
then
incubated
with
the
RNA
probes
at

242 65C
for
24
hours.
We
enriched
the
post-hybridization
libraries
using
TruSeq
adapter
primers
with

Phusion⇤ High-Fidelity
DNA
Polymerase
(New
England
Biolabs
Inc.)
for
22
cycles.
Following

PCR
enrichment
we
cleaned
the
libraries
with
AMPure
XP
beads.
Enriched
DNA
libraries
were
quantified
using
qPCR
(Applied
Biosystems
Inc.)
with
primers
targeting
five
loci
mapping
to
the
Anolis
carolinensis
genome.
Final
libraries
were
verified
using
the
Agilent
Tape-station
2200

(Agilent
Technologies).
The
final
sample
libraries
were
pooled
in
equimolar
ratios
and
sequenced
using
an
Illumina
HiSeq2500
at
the
DNAsc
(DNA
sequencing
center)
at
Brigham
Young
University

(BYU).

Bioinformatics
and
data
processing.
The
raw
DNA
sequence
reads
were
demultiplexed
based
on
unique
dual-indexed
sequence
tags
using
Casava
(Illumina).
We
pre-processing
the
raw
sequences
reads
with
a
custom
Rscript
(Illumina
prep
function
github:
XXXXX)
to
reformat
the
sequences
names
with
the
proper
sequence
adapters
to
make
them
compatible
input
files
for
PHYLUCE62
and
illumiprocessor63.
We
used
illumiprocessor
v
2.0.763
to
batch
process
the
demultiplexed
sequence
reads
and
trim
adapter
contamination
in
preparation
for
Trimmomatic64
to
remove
low-quality
reads,
adapter
sequences
and
low-quality
end
reads.
We
assembled
clean
reads
for
each
species
using
Trinity65
and
used
the
PHYLUCE
pipeline62
to
assemble
loci
across
species
(TableS1).
Mul- tiple
sequence
alignments
were
performed
for
each
locus
using
MAFFT66
and
PHYLUCE62
was
used
to
assemble
loci
across
species
and
long
ragged-ends
were
trimmed
to
reduce
missing
or
incomplete
data.
Phylogenomic
algorithms
for
both
concatenated
and
species
tree
approaches
pro- vide
the
highest
branch
support
values
when
including
loci
with
up
to
50%
of
the
missing
taxa28,
and
excluding
all
missing
data
from
the
dataset
might
be
a
problematic
guideline
for
setting
a
stan-

243 dard
of
missing
data.
We
assembled
three
different
datasets
(50p,
60p,
and
75p),
whereas
each
loci
per
dataset
must
contain
50%,
60%,
and
75%
of
the
taxa
respectively.
All
datasets
were
assembled
using
the
PHYLUCE62
pipeline
for
all
down
stream
analyses.

We
obtained
targeted
sequence
capture
data
from
541
UCE
and
44
SToL
loci
for
71
indi- viduals
(Table
S1)
with
three
species
that
were
not
successful
during
the
hybrid
enrichment
(C.
affinis,
C.
hangus,
C.
narathiawatensis)
and
were
removed
from
downstream
analyses.
The
lack
of
success
for
these
two
samples
could
be
caused
from
inefficiently
during
the
hybridization
step
or
low
sequencing
effort34.
We
combined
the
recombination
activating
protein
1
(RAG1)
from
the
SToL
capture
dataset
in
combination
with
the
RAG1
dataset
of
Grismer
et
al.
201426,
as
they
are
contiguous
sequences
and
allowed
us
to
incorporate
the
three
species
that
were
not
successful
during
the
hybrid
enrichment,
completing
a
dataset
of
all
inclusive
species
of
Cnempasis
where
genetic
material
is
available
(53
of
56).
Summary
data
for
the
541
UCE
and
the
44
SToL
loci
were
calculated
using
scripts
available
at
https://github.com/dportik/Alignment
Assessment67
(Figure

S3A–E).
Frequency
distributions
for
proportions
of
the
genomic
data
are
presented
on
a
per
locus
basis
for
69
of
the
71
individuals
sequenced
and
we
found
a
positive
correlation
between
the
num- ber
of
informative
sites
and
the
alignment
length
(R2=0.0854,
p<
0.0000,
Figure
S3F).
The
final
alignments
were
composed
21
outgroup
and
53
ingroup
taxa
for
514
loci.

Phylogenomic
analyses.
We
inferred
phylogenetic
relationships
within
Cnemaspis
using
unpar- titioned
Maximum
Likelihood
(ML),
Bayesian
Inference
(BI),
and
coalescent
based
species
tree
methods.
All
analyses
were
rooted
based
on
the
most
distant
relative
(Correlophus
cilliatus
and

Lialis
burtonis)
following
Gamble
et
al.57,
allowing
to
test
for
the
monophyly
of
Cnemaspis
with

244 respect to the other outgroup species. For the concatenated ML analyses we used the IQ-TREE software68 with the model of molecular evolution set to GTR+ and nodal support estimated with

1000 bootstrap pseudoreplicates via the ultrafast bootstrap approximation algorithm69. The BI analyses were estimated using ExaBayes70 based on four runs for 100,000 generations each, af- ter which convergence was assessed using the postProcParam command; we assumed stationarity if ESS values were 200, and computed a consensus tree using the consense command in Ex- aBayes. Nodal support for the concatenated analysis are posterior probabilities (PP) and IQ-TREE rapid bootstraps (BS), respectively, and nodes with (PP) and IQ-TREE rapid bootstraps (BS), re- spectively, and nodes with PP 0.95 and BS 90 are considered to be well-supported69, 71, 72.

We estimated a species tree from 514 UCE/sequence capture loci by first estimating gene trees for each locus using a search of 200 iterations for the best tree, and the resulting gene trees with the highest respective likelihood score were used as the input trees for downstream species tree analyses in ASTRAL-II (A-II)73. In addition to the initial branch support values provided from the species tree, which is considered to be more reliable74, we estimated nodal support using

100 bootstrap replicates for each gene generated in RAxML v7.5.575, and then used the default site-only resampling76 in A-II73, which is based on the quadripartitions and not the more often used branch bipartitions74. To further investigate specific nodes/branches in the deeper nodes for the Cnemaspis nodes we calculated quartet support values in A-II73. The quartet support values are the posterior estimate at a given branch where each genetree quartet agrees with the respective branch.

To estimate a species tree without first estimating gene trees, we used the program SVDQuar-

245 tets
(SVDQ)77,
as
implemented
in
a
test
version
of
Paup*v4.0a15078.
This
algorithm
randomly
samples
quartets
using
a
coalescent
model
and
and
a
quartet
amalgamation
heuristic
to
gener- ate
a
species
tree,
and
has
proven
useful
and
accurate
for
estimating
species
trees
from
complete
alignments
of
large
genomic
datasets79.
We
evaluated
all
possible
quartet
scores
from
the
entire
alignment
using
the
multispecies
coalescent
tree
model,
and
performed
100
bootstrap
replicates
to
calculate
nodal
support.
The
program
Quartet
MaxCut
v2.1.080
was
used
to
construct
a
species
tree
from
the
sampled
quartets.

To
estimate
times
of
divergence
within
the
genus
Cnemaspis
and
their
relatives
we
used
a
constrained
concatenated
ExaBayes
UCE
tree
based
on
514
loci
(267,912
bp)
as
the
guide
tree
in
MCMCtree
in
PAML
v4.981.
We
used
one
fossil
calibration
and
a
root
height
for
the
crown
gekkotans.
The
fossil
calibration
was
based
on
the
divergence
between
Sphaerodactylus
roosevelti
and
S.
townsendi
as
estimated
from
an
amber-preserved
fossil
Sphaerodactylus
from
Hispanola
dated
15–20
million
years
ago
(MYA)82.
We
set
the
root
height
of
crown
gekkotans
with
an
upper
bound
of
125
MYA
and
a
lower
bound
of
95
my
based
on
previously
used
constraints30,83–86.
The
divergence
times
were
estimate
using
the
program
MCMCtree
in
PAML
v4.981.
Divergence
times
were
estimated
with
the
independent
rates
model
and
the
Birth-Death
process
for
species
sampling.

The
size
of
our
data
set
precluded
implementation
of
a
full
Markov
chain
Monte
Carlo

(MCMC)
iteration,
so
we
used
an
approximate
likelihood
algorithm87,88.
We
first
calculated
maximum-likelihood
estimates
for
the
branch
lengths,
a
gradient
vector
and
Hessian
matrix,
using
the
internal
programs
BaseML
and
CodeML
in
PAML
v4.981.
The
most
complex
model
available
in
MCMCtree
was
applied
(HKY+)
with
four
categories,
which
allows
for
rate
variation
among

246 nucleotides.
The
second
step
in
our
analysis
used
the
MCMC
algorithm
to
estimate
the
divergence

times
on
the
given
guide
tree,
and
the
estimated
gradient
vectors
along
with
the
Hessian
matrices

were
used
to
generate
the
Taylor
expansion
for
the
log-likelihoods89.
The
incorporation
of
these

approximate
likelihood
estimates
on
large
datasets
decreases
the
computational
time
and
makes

these
analyses
tractable.
These
analyses
were
run
for
106
generations,
sampling
every
103
gen-

erations,
and
with
a
10%
burnin.
The
MCMC
outputs
were
loaded
into
Tracer
v1.690
to
assess convergence,
and
stationarity
was
assumed
when
effective
sample
sizes
were

200.

Diversification
and
ancestral
area
reconstructions.
To
test
for
rapid
rates
of
diversification
in

Cnemaspis,
and
to
estimate
speciation
and
extinction
rates,
we
used
the
Bayesian
Analysis
of

Macroevolutionary
Mixtures
(BAMM)
program
v2.531,
which
detects
and
quantifies
rate
hetero-

geneity
along
the
branches
of
a
phylogenetic
tree.
We
analyzed
our
complete
dataset
(ingroup
+

outgroup
terminals),
and
then
a
Cnemaspis-only
dataset
separately,
to
be
sure
that
younger
rate
shifts
were
not
being
obscured
by
the
inclusion
of
the
much
older
outgroups.
All
BAMM
analyses

were
run
for
106
generations
and
event
data
were
sampled
every
103
generations,
and
we
simu- lated
the
prior
distribution
of
the
number
of
rate
shifts
using
default
priors.
BAMM
accommodates

unsampled
taxa
by
incorporating
a
nonrandom
incomplete
taxon
sampling
correction
into
the
like-

lihood
equation.
We
discarded
10%
of
the
generations
as
burnin
after
assessing
convergence
in

BAMMtools31,91,92.
We
checked
effective
sample
sizes
using
the
CODA
package93
and
assumed

convergence
if
ESS
>300.
The
rate
shifts,
speciation
rates,
extinction
rates,
and
rate-through-

time
curves
were
summarized
and
visualized
using
the
R
v3.1.294
package
BAMMtools92
and
all

247 speciation
and
extinction
rates
are
mean
rates
and
not
constant
through
time.

Independently
of
BAMM
we
plotted
fluctuation
rates
of
a
dated
phylogeny
and
visualized
the
diversification
history
of
Cnemaspis
via
construction
of
a
lineage-through-time
(LTT)
plot.

LTT
plots
and
their
associated
statistics
have
been
used
for
testing
simultaneous
radiations
versus
constant
rates
of
speciation95,96.
We
constructed
a
LTT
plot
from
the
time-calibrated
tree
generated
in
MCMCtree
using
the
R
package
ape
v4.097,
and
tested
for
a
significant
departure
from
the
null
hypothesis
(a
constant
rate
of
cladogenesis
in
Cnemaspis)
using
a
Monte
Carlo
constant
rates
test

(MCCRT98,99)
of
diversification,
as
implemented
in
the
R
package
Laser
v2.4-1100.
The
net
diver- sification
rate
statistic
()
was
estimated
for
the
time-calibrated
MCMCtree
using
Ape
v4.0.
To
account
for
missing
taxa
we
simulated
20,000
constant
rate
pure-birth
trees
with
random
pruning
to
mimic
incomplete
sampling,
until
the
empirical
number
of
taxa
was
reached
(n
=
53[55]).
The
estimated-statistic
was
used
to
compare
the
relative
positions
of
the
nodes
in
the
phylogeny
to
the
expected
distribution
under
a
constant
rate
of
speciation
model
of
diversification.
A
negatively
significant-statistic
<-1.64598
is
expected
to
be
a
good
measure
of
a
rapid
radiation.

To
estimate
ancestral
areas
at
nodes
of
the
Cnemaspis
time
tree,
we
first
pruned
the
tree
to
in- clude
only
Cnemaspis,
and
we
then
designated
biogeographic
regions
throughout
the
distribution
of
Cnemaspis
across
Southeast
Asia.
We
designated
13
biogeographic
regions
(Figure
1B)
that
either:
(1)
correspond
to
all
of
the
islands
in
a
given
archipelago;
(2)
were
previously
designated
regions
from
the
literature
(see
below);
or
(3)
were
previously
recovered
clades
of
Cnemaspis
and
do
not
correspond
to
any
political
boundaries.
This
last
group
includes:
(A)
Pulau
Bidong;
(B)

Seribuat
Archipelago,
Malaysia;
(C)
Ca
Mau
Islands,
Vietnam;
(D)
Borneo;
(E)
Sumatra,
Indone-

248 sia;
(F)
Peninsular
Malaysia
(from
the
southern
portion
of
the
Kangar-Pattani
line
south
including

Singapore);
(G)
Pattani
(from
the
northern
border
of
the
Kangar-Pattani
line
north
to
the
south- ern
part
of
the
Isthmus
of
Kra);
(H)
Central
Indochina
(from
the
northern
border
of
the
Isthmus
of
Kra
east
of
the
Salween
River,
and
west
of
the
Mekong
River,
and
the
greater
Mekong
Delta
region47);
(I)
Langkawi
Archipelago,
Malaysia;
(J)
Natuna
Archipelago,
Indonesia;
(K)
Perhentian

Archipelago,
Malaysia;
(L)
Pulau
Pinang,
Malaysia;
and
(M)
Hon
Tre
Island,
Vietnam.

To
infer
the
ancestral
ranges
of
Cnemaspis
we
used
a
probabilistic-statistical
modeling
ap- proach
in
the
R94
package
BioGeoBEARS101,
to
test
alternative
diversification.
BioGeoBEARS
uses
the
Akaike
information
criteria
(AIC)
in
a
statistical
model
fitting
approach
to
select
the
best

fit
model
for
ancestral
geographic
range
reconstructions
for
the
data.
We
tested
four
models
in

BioGeoBEARS,
including:
(1)
a
likelihood
version
of
the
maximum
parsimony
model
Dispersal-

Vicariance
Analysis
DIVA
(‘DVIALIKE’)
model102);
(2)
the
Dispersal-Extinction
Cladogenesis

(DEC)
model103,104
which
has
a
common
conceptual
frame
work
with
traditional
with
the
DIVA
model
of
ancestral
character
reconstructions,
in
which
biogeographic
areas
are
coded
as
discrete
characters
and
species’
ranges
are
coded
as
binary
presence/absence
data,
and
the
ranges
evolve
along
the
phylogenetic
branches
by
lineage
movement
(dispersal
from
one
area
to
another)
and
local
extinction105;
and
(3)
the
Bayesian-based
BayArea
(‘BAYAREALIKE’)
model106
which
ex- tends
the
application
of
more
realistic
problems
involving
large
numbers
of
biogeographic
areas
by
marginalizing
over
all
possible
biogeographic
histories106.

In
combination
with
these
models
we
also
incorporated
a
jump-dispersal
or
founder
event

speciation
parameter
(+J),
was
incorporated;
this
step
has
been
shown
to
be
useful
for
datasets
of

249 insular clades107. We used a discrete character presence/absence matrix for the above 13 biogeo- graphic regions for each species, based on specimen-verified locality data in Grismer et al. 201426 and Wood et al. 201725. We used likelihood ratio tests of the null (without +J) versus the alternative models (with +J) to see if we could reject scenarios without founder event speciation. All analyses were unconstrained, allowing each individual to have the same probability of colonizing among all adjacent areas or biogeographic regions. The maximum number of areas was set to three, with only four species occupying more than one biogeographic region with none of them occupying more than two.

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263 Acknowledgements
We
like
to
thank
Leonard
Jones,
Itzue
Caviedes-Solis,
Nassima
Bouzid,
and
Matt

McElroy,
for
mentorship
and
emotional
support
generating
the
genomic
data.
Juan
C.
Santos
for
assistance
with
R
code
and
general
discussions
on
comparative
methods.
For
discussions
and
advice
for
implementing

some
of
the
genomic
analyses
we
are
grateful
for
Alana
Alexandra
and
Steve
Levitt
that
greatly
improve
the
quality
of
the
manuscript.
I
would
like
to
thank
Phillip
Skipwith,
Daniel
Portik
and
Mario
dos
Reis
for
multiple
discussions
relating
to
divergence
time
estimates
using
large
genomic
datasets.
Partial
funding
was
supported
by
the
NSF
grant
EF-1241885
issued
to
JWS,
and
the
Department
of
Biology
at
Brigham

Young
University.
Funding
for
generation
of
the
molecular
data
was
supported
by
a
Doctoral
Dissertation

Improvement
Grant
(DDIG)
issued
to
JWSJ
and
PLWJ
(NSF
#1501198).
Partnerships
for
Enhanced
En- gagement
in
Research
(PEER)
Science
program
(grant
PGA-2000003545),
which
is
a
partnership
between
the
U.S.
Agency
for
International
Development
(USAID)
and
the
National
Science
Foundation
(NSF),
pro- vided
funding
for
AA
for
data
collection
on
this
project.

Competing
Interests
The
authors
declare
that
they
have
no
competing
financial
interests.

Correspondence
Correspondence
and
requests
for
materials
should
be
addressed
to
P.L.W.J.
(email:
[email protected]).

264 abe SH,Uiest an aasaHreooia olcina h nvriiSisMlyi,Pnn,Mlyi;UA,Uiest fTxsa Austin at Texas of University Thailand. UTAF, University, Kasetsart Malaysia; Museum Penang, Zoological Malaysia, R, Sains Museum, ZMKU Universiti series; Science field the Life at Bean Collection L. Herpetological Tony Malaysia Monte TG, Indonesia; Sains BYU, Java, Universiti Cibinung, follows: USMHC, Bogoriense, Gamble; Zoologicum as Museum are MZB, abbreviations Collection; Herpetological number University Voucher Sierra La LSUHC, dataset. University; Young capture Brigham UCE/sequence the for used Specimens S1: Table Supplemental Material nmsi flavolineata Cnemaspis flavigaster Cnemaspis chanthaburiensis Cnemaspis chanardi Cnemaspis caudanivea Cnemaspis boulengerii Cnemaspis biocellata Cnemaspis bidongensis Cnemaspis bayuensis Cnemaspis baueri Cnemaspis aurantiacopes Cnemaspis argus Cnemaspis Samples SH 0917 52032043824 9319310 9 301 19 2841 332.0742358 456270 1374 405 8079 LSUHC 8835 LSUHC 6 9338 LSUHC 399 9567 LSUHC 8577 LSUHC 42 9278 LSUHC 8927 8789 LSUHC 400.5880994 11445 LSUHC 1386836 9072 LSUHC 3462 7302 LSUHC 10835 LSUHC 8612 LSUHC Vouchers otg oa plnt length length bp Total Contigs 70228739164595 5 7 2492 258 22 13 486 470 280 245 26 374 8 156 460 18 272 150 18 9156 460 3348 389.1565445 356 49 2 299.346247 21 14 2229867 12791 419 148 439 421.7579948 4959 741780 32 530 5730 309.1884146 21 1463922 2580 30 266 2478 535 4 398 389.9130932 507069 500 15381 3471 18 1726 1363916 334.0394046 1640 305.2952586 253 248 6934324 73 16786 3498 283314 368.4781356 15591 20759 8 462.4124224 4474430 928 1408 1116726 12143 303.4330669 2415 607473 2002 enMxCnisUESo Total SToL UCE Contigs Max Mean > k oilc loci loci loci 1kb

265 al 1 . . S1. Table nmsi paripari Cnemaspis omari Cnemaspis nuicamensis Cnemaspis niyomwanae Cnemaspis nigridia Cnemaspis neangthyi Cnemaspis mumpuniae Cnemaspis monachorum Cnemaspis mcguirei Cnemaspis mahsurae Cnemaspis lineogularis Cnemaspis limi Cnemaspis kumpoli Cnemaspis kendallii Cnemaspis karsticola Cnemaspis huaseesom Cnemaspis harimau Cnemaspis grismeri Cnemaspis SH 9186 LSUHC 9978 LSUHC 8649 LSUHC 9568 LSUHC 9170 LSUHC 8515 LSUHC 10166 MZBLace 10807 LSUHC 8853 LSUHC 11828 LSUHC 00728 R ZMKU SH 3902 LSUHC 8848 LSUHC 9178 LSUHC 9055 LSUHC 9458 LSUHC 9667 LSUHC 9733 LSUHC 5327 6784 7264 7072 10096 4421 35748 3132 10527 1556 5522 2027 17075 4517 14910 3493 4113 1267 1931742 2677116 2643370 2628614 3230439 1575057 11306444 1164791 3566139 712900 2032636 960099 5431575 1524574 4727367 1323571 1544414 386531 362.6322508 394.6220519 363.9000551 371.6931561 319.9721672 356.2671341 316.2818619 371.9000639 338.7611855 458.1619537 368.0977907 473.6551554 318.1010249 337.5191499 317.060161 378.9209848 375.4957452 305.0757695 13851 6463 14478 15750 11559 2766 16474 8766 13705 4105 15801 13894 16943 12950 15629 8723 12675 2013 68 163 104 151 82 61 447 51 98 42 80 38 119 39 100 48 43 10 485 369 480 479 405 374 489 504 419 465 341 449 499 486 464 391 467 189 25 19 30 32 15 18 31 30 22 26 16 31 26 22 25 14 26 7 510 388 510 511 420 392 520 534 441 491 357 480 525 508 489 405 493 196

266 al 1 . . S1. Table nmsi temiah Cnemaspis sundainsula Cnemaspis stongensis Cnemaspis thachanaensis Cnemaspis Cnemaspis Cnemaspis siamensis Cnemaspis shahruli Cnemaspis selamatkanmerapoh Cnemaspis roticanai Cnemaspis rajabasa Cnemaspis punctatonuchalis Cnemaspis psychedelica Cnemaspis pseudomcguirei Cnemaspis phangngaensis Cnemaspis perhentianensis Cnemaspis peninsularis Cnemaspis pemanggilensis Cnemaspis p o.2 nov. sp. 1 nov. sp. SH 9817 LSUHC 10156 MZBLace 11089 LSUHC 00729 R ZMKU SH 9570 LSUHC 9569 LSUHC 9474 LSUHC 9613 LSUHC 11015 LSUHC 11815 LSUHC 7766 ENS 12453 LSUHC 9244 LSUHC 9145 LSUHC 62537 BYU 8700 LSUHC 0228 USMHC 8014 LSUHC 857 335 2445 4887 7034 646 289 1158 2848 2484 3978 8015 5401 11139 1909 7232 16150 4614 281424 99276 801773 1739130 86914 479578 861428 1076916 1171332 2499492 1561206 3349907 735157 2104491 5426317 1397381 2409882 249515 328.3827305 296.3462687 327.9235174 355.8686311 342.6047768 386.24613 300.7404844 414.1433506 302.4676966 433.5410628 294.4524887 311.8517779 289.0586928 300.7367807 385.1005762 290.9970962 335.9948607 302.8567404 3665 864 5654 6503 1323 12904 4031 4918 7678 7981 10028 12114 4799 14284 13227 3369 6942 645 6 0 33 97 66 0 1 9 7 47 14 79 12 74 18 38 293 35 239 124 156 269 368 483 85 369 224 420 170 249 338 309 179 228 501 184 6 5 3 13 3 23 8 26 10 13 13 16 17 10 30 7 9 20 245 129 159 282 88 392 232 446 180 262 351 325 196 238 531 191 377 503

267 al 1 . . S1. Table pardcyu roosevelti Sphaerodactylus macrolepis Sphaerodactylus wirshingii Phyllodactylus laticauda Phelsuma picta Paroedura burtonis Lialis turcicus Hemidactylus mabouia Hemidactylus vittatus Gonatodes ocellatus Gonatodes mutilata Gehyra scaber Cyrtopodion ciliatus Correlophus variegatus Coleonyx persica Agamura Outgroups vandeventeri Cnemaspis tucdupensis Cnemaspis SH 12328 LSUHC TG2384 TG2147 TG2385 TG1616 TG2139 TG2114 TG2125 TG2142 TG1845 TG1855 TG0109 TG2314 TG0179 TG0188 SH 12449 LSUHC 8631 LSUHC 4257 12546 7866 7764 8832 12806 340 6004 12256 20639 6433 24800 1105 19048 60658 1327 913 1284436 4171741 2881395 2879736 3349236 4709784 102614 1752371 4476060 6797555 1918054 7740463 465188 6926627 18572940 677694 271781 301.7232793 332.5156225 366.3100687 370.9088099 379.2160326 367.7794784 301.8058824 291.8672552 365.2137728 329.3548622 298.1585574 312.1154435 420.9846154 363.6406447 306.1911042 510.6963075 297.67908 3209 11606 4116 16201 10226 15493 1125 6241 16771 15735 10116 15488 3170 16872 7486 8817 2228 25 127 131 202 215 539 4 24 180 278 37 108 10 495 512 12 6 273 471 286 416 305 311 82 207 299 416 285 387 345 322 458 435 35 10 32 7 24 13 14 0 11 12 17 12 26 18 12 27 22 1 283 503 293 440 318 325 82 218 311 433 297 413 363 334 485 457 36

268 al 1 . . S1. Table Tropiocolotes rapicauda Thecadactylus przewalskii Teratoscincus keyserlingii Teratoscincus Stenodactylus townsendi Sphaerodactylus cf. cf. tripilotanus sthenodactylus TG2354 TG1864 TG233 TG2382 TG0185 TG2150 1823 32594 16085 3529 6393 23647 756570 10374532 5652107 1552260 2236146 7673801 415.0137137 318.29576 351.3899285 439.8583168 349.7803848 324.5147799 4172 16664 15606 8079 10310 22498 37 351 287 117 73 387 462 401 498 466 391 490 28 14 34 31 24 31 490 415 532 497 415 521

269 Table S2: The results of the model fitting from the ancestral area reconstructions in Bio-

GeoBears. d= is the rate of dispersal or range expansion, e= is the extinction rate, and j= is the jump-dispersal rate.

LnL numparams d e j AICc AICc wt

DEC -123.2 2 0.19 0.73 0 250.7 1.0e-09

DEC+J -108.1 3 0.062 0.72 0.018 222.7 0.0012

DIVALIKE -114.4 2 0.20 1.0e-12 0 233.1 6.7e-06

DIVALIKE+J -101.4 3 0.061 1.0e-12 0.016 209.3 1.00

BAYAREALIKE -146.6 2 0.20 2.92 0 297.4 7.2e-20

BAYAREALIKE+J -122.4 3 0.047 2.94 0.017 251.4 7.2e-10

270 Mau C C psychedelica Ca A. C C 12.1 C C boulengerii j B B limi D D D D paripari Southern Sunda Outgroups 21.8 D D 5.9 D nigridia D D D 23.3 j J J sundainsula D D D D kendallii j 16.3 E E rajabasa v D D B B 7.6 13.2 B CDF B pemanggilensis 61.1 B B 10 B B baueri B B B BF peninsularis 6.9 A j B B A bidongensis 4.0 J J j J mumpuniae Out of the 3.1 J Sunda Shelf F F argus F F F F flavigaster Rapid radiation 16.4 v F F j K K perhentianensis DF 12.4 F F DF 4.8 F karsticola 41.9 F Malaysian Peninsula F F F F pseudomcguirei 24.8 B. 11.6 F FL shahruli 6.7 F F grismeri F F F F F F F F mcguirei H 14.8 F F j L L affinis 10.4 F F I j 5.2 I mahsurae F F F F 12.4 3.2 F F harimau M 9.3 F F temiah F F C F K F flavolineata F F F narathiwatensis G 11.3 F F F F F F F selamatkanmerapoh A j 9.0 I 39.2 F F F J 6.9 F hangus F F F B 5.9 F stongensis F F F F 3.4 F bayuensis I I sp. nov. 1 I L I Thai Peninsula Indochina Pattani 22.1 10.6 I I monachorum D I I j G G biocellata G G G 11.2 G niyomwanae E G G 5.9 G G kumpoli j H H lineogularis H H M M caudanivea H H 20.5 j C. 37.3 H H H H chanthaburiensis 15.0 Volcanoes H H H H neangthyi Highlands 11.6 H H H H aurantiacopes Land 9.6 H H H Lakes, Freshwater H tucdupensis H H Lakes, Brackish H H H nuicamensis Shallow Sea 33.4 5.4 H Carbonate Platforms H H punctatonuchalis Deep Sea H H H H huaseesom 60 Ma 40 Ma Trenches 18.7 Paleocene Late Eocene H H H GH siamensis 14.0 H H H H vandeventeri 11.8 H H H H H thachanaensis 26.6 7.2 H H 6.7 H phangngaensis H H H GH chanardi H H 8.7 I I sp. nov. 2 j I I I I roticanai 5.7 10 Ma I 30 Ma 20 Ma G G omari Mid Oligocene Early Miocene Late Miocene j 5.0 Upper/Late Paleocene Eocene Oligocene Miocene Pli. P 80 60 40 20 0 MYA

Figure 1: A. Chronogram of Cnemaspis based on a concatenated ExaBayes analysis of 514-locus guide tree (full tree in supplemental Figure S5). Numbers near the nodes are mean divergence time estimates from fossil and root height calibrations implemented in MCMCtree. The topology and all nodal support values are identical to Figure S4. Unconstrained ancestral area reconstructions inferred from the DIVALIKE+J model in BioGeoBears [1]. The black star indicates the rate-shift detected from the BAMM analysis (Probability=70%). B. Biogeographic regions for the ancestral range reconstructions (abbreviations described in the materials and methods). Arrows indicate a dispersal or vicariant event from one biogeographic region to another, j = jump-dispersal or founder event and v=vicariant event. C. Paleogeographic reconstructions are modified from Hall[2, 3]. 271 Figure S1: Distribution of the genus Cnemaspis based on Grismer et al. 2014[4]; colors identify major clades. The black circle is C. laoensis and its placement into one of the major clades is unknown due to the lack of genetic material. The map is generated using a global one arc-minute grid obtained from the General Bathymetric Chart of the Oceans (GEBCO).

272 A. IQTREE 50p B. IQTREE 60p B. IQTREE 75p

100 Correlophus ciliatus TG2314 100 Lialis burtonis TG2114 100 Lialis burtonis TG2114 Lialis burtonis TG2114 Correlophus ciliatus TG2314 Correlophus ciliatus TG2314 Coleonyx variegatus TG0179 Coleonyx variegatus TG0179 Coleonyx variegatus TG0179 100 Teratoscincus przewalskii TG233 100 Teratoscincus przewalskii TG233 100 Teratoscincus przewalskii TG233 Teratoscincus keyserlingii TG2382 Teratoscincus keyserlingii TG2382 Teratoscincus keyserlingii TG2382 100 Gonatodes ocellatus TG1855 100 Gonatodes vittatus TG1845 100 Gonatodes vittatus TG1845 100 100 100 100 100 100 Gonatodes vittatus TG1845 100 100 Gonatodes ocellatus TG1855 100 Gonatodes ocellatus TG1855 Sphaerodactylus macrolepis TG2147 Sphaerodactylus macrolepis TG2147 Sphaerodactylus macrolepis TG2147 100 100 100 Sphaerodactylus townsendi TG2150 Sphaerodactylus townsendi TG2150 98 Sphaerodactylus townsendi TG2150 100 100 100 Sphaerodactylus roosevelti TG2384 100 Sphaerodactylus roosevelti TG2384 100 Sphaerodactylus roosevelti TG2384 100 Phyllodactylus wirshingii TG2385 100 Thecadactylus rapicauda TG1864 100 Thecadactylus rapicauda TG1864 Thecadactylus rapicauda TG1864 Phyllodactylus wirshingii TG2385 Phyllodactylus wirshingii TG2385 Gehyra mutilata LSUHC12328 Gehyra mutilata LSUHC12328 Gehyra mutilata LSUHC12328 53 100 100 Hemidactylus turcicus TG2125 80 100 Hemidactylus mabouia TG2142 Paroedura picta TG2139 100 Hemidactylus mabouia TG2142 100 Hemidactylus turcicus TG2125 100 59 Phelsuma laticauda TG1616 100 100 100 Tropiocolotes cf tripilotanus TG2354 100 Tropiocolotes cf tripilotanus TG2354 100 Hemidactylus mabouia TG2142 100 Stenodactylus cf sthenodactylus TG0185 100 Stenodactylus cf sthenodactylus TG0185 Hemidactylus turcicus TG2125 Agamura persica TG0188 100 71 Cyrtopodion scaber TG0109 57 100 30 100 Tropiocolotes cf tripilotanus TG2354 Agamura persica TG0188 Cyrtopodion scaber TG0109 100 Stenodactylus cf sthenodactylus TG0185 Paroedura picta TG2139 Paroedura picta TG2139 Cyrtopodion scaber TG0109 Phelsuma laticauda TG1616 Phelsuma laticauda TG1616 Agamura persica TG0188 100 Cnemaspis psychedelica LSUHC 9244 100 Cnemaspis boulengerii LSUHC 9278 100 Cnemaspis psychedelica LSUHC 9244 61 Cnemaspis boulengerii LSUHC 9278 99 Cnemaspis psychedelica LSUHC 9244 Cnemaspis boulengerii LSUHC 9278 Cnemaspis limi LSUHC 3902 Cnemaspis limi LSUHC 3902 100 Cnemaspis limi LSUHC 3902 86 90 88 100 Cnemaspis paripari LSUHC 9186 100 Cnemaspis paripari LSUHC 9186 100 Cnemaspis nigridia LSUHC 9170 98 100 Cnemaspis nigridia LSUHC 9170 100 100 Cnemaspis nigridia LSUHC 9170 100 Cnemaspis paripari LSUHC 9186 Cnemaspis sundainsula MZBLace 10156 Cnemaspis sundainsula MZBLace 10156 Cnemaspis sundainsula MZBLace 10156 Cnemaspis kendallii LSUHC 9178 Cnemaspis kendallii LSUHC 9178 Cnemaspis pemanggilensis LSUHC 8014 100 100 98 Cnemaspis rajabasa UTAF14 ES ENS7766 Cnemaspis rajabasa UTAF14 ES ENS7766 100 Cnemaspis kendallii LSUHC 9178 84 93 100 100 100 Cnemaspis pemanggilensis LSUHC 8014 100 Cnemaspis pemanggilensis LSUHC 8014 Cnemaspis baueri LSUHC 7302 91 6799 100 Cnemaspis baueri LSUHC 7302 63 Cnemaspis baueri LSUHC 7302 Cnemaspis peninsularis HC 0228a 94 Cnemaspis peninsularis HC 0228a 98 Cnemaspis peninsularis HC 0228a 96 Cnemaspis mumpuniae MZBLace 10166 GL115 95 Cnemaspis bidongensis LSUHC 11445 100 Cnemaspis bidongensis LSUHC 11445 96 Cnemaspis bidongensis LSUHC 11445 99 Cnemaspis mumpuniae MZBLace 10166 GL115 99 Cnemaspis mumpuniae MZBLace 10166 GL115 84 Cnemaspis rajabasa UTAF14 ES ENS7766 Cnemaspis argus LSUHC 10835 Cnemaspis argus LSUHC 10835 100 Cnemaspis sp 1 LSUHC 9569 x MS 286 100 100 100 Cnemaspis flavigaster LSUHC 8835 Cnemaspis flavigaster LSUHC 8835 100 Cnemaspis monachorum LSUHC 10807 Cnemaspis perhentianensis LSUHC 8700 Cnemaspis perhentianensis LSUHC 8700 Cnemaspis biocellata LSUHC 8789 100 78 100 86100 100 100 Cnemaspis karsticola LSUHC 9055 Cnemaspis karsticola LSUHC 9055 Cnemaspis niyomwanae LSUHC 9568 100 100 Cnemaspis pseudomcguirei LSUHC 9145 Cnemaspis pseudomcguirei LSUHC 9145 100 Cnemaspis kumpoli LSUHC 8848 Cnemaspis shahruli LSUHC 9163 Cnemaspis mcguirei LSUHC 8853 Cnemaspis flavigaster LSUHC 8835 70 97 82 100 Cnemaspis grismeri LSUHC 9733 10067 Cnemaspis grismeri LSUHC 9733 100 Cnemaspis perhentianensis LSUHC 8700 100 Cnemaspis mcguirei LSUHC 8853 Cnemaspis affinis LSUHC 6759 Cnemaspis karsticola LSUHC 9055 89 96 Cnemaspis affinis LSUHC 6759 100 Cnemaspis mahsurae LSUHC11828 Cnemaspis flavolineata LSUHC 8079 Cnemaspis mahsurae LSUHC11828 91 Cnemaspis harimau LSUHC 9667 Cnemaspis shahruli LSUHC 9163 100 64 99 87 Cnemaspis harimau LSUHC 9667 Cnemaspis shahruli LSUHC 9163 67 Cnemaspis narathiwatensis USMHC 1350 55 100 93 Cnemaspis temiah LSUHC 9817 88 Cnemaspis temiah LSUHC 9817 98 Cnemaspis selamatkanmerapoh LSUHC 11015 58 Cnemaspis flavolineata LSUHC 8079 Cnemaspis flavolineata LSUHC 8079 93 Cnemaspis hangus HC 225 95 Cnemaspis narathiwatensis USMHC 1350 90 Cnemaspis narathiwatensis USMHC 1350 58 Cnemaspis stongensis LSUHC 11089 75 Cnemaspis selamatkanmerapoh LSUHC 11015 Cnemaspis hangus HC 225 93 Cnemaspis bayuensis LSUHC 9072 100 85 98 87 96 Cnemaspis hangus HC 225 95 Cnemaspis selamatkanmerapoh LSUHC 11015 Cnemaspis temiah LSUHC 9817 52 Cnemaspis stongensis LSUHC 11089 44 Cnemaspis stongensis LSUHC 11089 Cnemaspis pseudomcguirei LSUHC 9145 48 99 Cnemaspis bayuensis LSUHC 9072 99 Cnemaspis bayuensis LSUHC 9072 82 Cnemaspis mcguirei LSUHC 8853 77 100 Cnemaspis sp 1 LSUHC 9569 x MS 286 100 Cnemaspis sp 1 LSUHC 9569 x MS 286 Cnemaspis grismeri LSUHC 9733 Cnemaspis monachorum LSUHC 10807 Cnemaspis monachorum LSUHC 10807 50 83 Cnemaspis affinis LSUHC 6759 100 100 97 Cnemaspis biocellata LSUHC 8789 Cnemaspis biocellata LSUHC 8789 Cnemaspis mahsurae LSUHC11828 100 100 Cnemaspis niyomwanae LSUHC 9568 Cnemaspis niyomwanae LSUHC 9568 90 Cnemaspis harimau LSUHC 9667 100 Cnemaspis kumpoli LSUHC 8848 100 Cnemaspis kumpoli LSUHC 8848 Cnemaspis aurantiacopes LSUHC 8612 Cnemaspis sp nov 3 LSUHC 12480 Cnemaspis sp nov 3 LSUHC 12480 71 Cnemaspis sp nov 3 LSUHC 12480 100 Cnemaspis caudanivea LSUHC 8577 100 Cnemaspis caudanivea LSUHC 8577 100 Cnemaspis caudanivea LSUHC 8577 100 100 Cnemaspis chanthaburiensis LSUHC 9338 Cnemaspis chanthaburiensis LSUHC 9338 87 Cnemaspis neangthyi LSUHC 8515 99 100 82 100 Cnemaspis neangthyi LSUHC 8515 88 Cnemaspis neangthyi LSUHC 8515 99 Cnemaspis chanthaburiensis LSUHC 9338 78 Cnemaspis aurantiacopes LSUHC 8612 98 Cnemaspis aurantiacopes LSUHC 8612 Cnemaspis argus LSUHC 10835 94 Cnemaspis tucdupensis LSUHC 8631 Cnemaspis tucdupensis LSUHC 8631 99 Cnemaspis tucdupensis LSUHC 8631 100 100 71 98 100 Cnemaspis nuicamensis LSUHC 8649 100 Cnemaspis nuicamensis LSUHC 8649 Cnemaspis nuicamensis LSUHC 8649 Cnemaspis punctatonuchalis LSUHC 12453 Cnemaspis punctatonuchalis LSUHC 12453 Cnemaspis siamensis LSUHC 9474 100 100 Cnemaspis huaseesom LSUHC 9458 Cnemaspis huaseesom LSUHC 9458 100 100 Cnemaspis vanderventeri LSUHC12449 85 60 Cnemaspis siamensis LSUHC 9474 Cnemaspis siamensis LSUHC 9474 Cnemaspis sp nov 2 LSUHC 12470 54 Cnemaspis vanderventeri LSUHC12449 94 Cnemaspis sp nov 2 LSUHC 12470 96 Cnemaspis punctatonuchalis LSUHC 12453 100 100 100 100 Cnemaspis sp nov 2 LSUHC 12470 100 Cnemaspis vanderventeri LSUHC12449 99 Cnemaspis huaseesom LSUHC 9458 97 Cnemaspis sp nov 1 LSUHC 12438 98 Cnemaspis sp nov 1 LSUHC 12438 98 Cnemaspis sp nov 1 LSUHC 12438 100 Cnemaspis chanardi LSUHC 9567 100 Cnemaspis chanardi LSUHC 9567 100 Cnemaspis chanardi LSUHC 9567 Cnemaspis sp 2 LSUHC 9570 x MS 283 Cnemaspis sp 2 LSUHC 9570 x MS 283 Cnemaspis roticanai LSUHC11815 100 Cnemaspis roticanai LSUHC11815 100 Cnemaspis roticanai LSUHC11815 100 Cnemaspis sp 2 LSUHC 9570 x MS 283 0.01 94 Cnemaspis omari LSUHC 9978 0.01 92 Cnemaspis omari LSUHC 9978 0.005 98 Cnemaspis omari LSUHC 9978

Figure S2: Maximum Likelihood trees estimated in IQTREE[5, 6] using three di↵erent sized datasets: A. 50p (267,912 bp/514 loci), B. 60p (231,027 bp/440 loci), C.75p (57,940bp/105 loci).

273 A. Concatenated UCEs B. ASTRAL−II species tree Correlophus ciliatus TG2314 Correlophus ciliatus TG2314 Lialis burtonis TG2114 Lialis burtonis TG2114 Coleonyx variegatus TG0179 Coleonyx variegatus TG0179 Teratoscincus przewalskii TG233 Teratoscincus przewalskii TG233 Teratoscincus keyserlingii TG2382 Teratoscincus keyserlingii TG2382 Gonatodes vittatus TG1845 Gonatodes vittatus TG1845 Gonatodes ocellatus TG1855 Gonatodes ocellatus TG1855 Sphaerodactylus macrolepis TG2147 Sphaerodactylus macrolepis TG2147 Sphaerodactylus townsendi TG2150 Sphaerodactylus townsendi TG2150 Sphaerodactylus roosevelti TG2384 0.84/86/100 Sphaerodactylus roosevelti TG2384 Thecadactylus rapicauda TG1864 Thecadactylus rapicauda TG1864 Phyllodactylus wirshingii TG2385 0.97/54/100 Phyllodactylus wirshingii TG2385 Gehyra mutilata LSUHC 12328 1.0/61/- Gehyra mutilata LSUHC 12328 Hemidactylus turcicus TG2125 Hemidactylus turcicus TG2125 Hemidactylus mabouia TG2142 Hemidactylus mabouia TG2142 Tropiocolotes cf. tripilotanus TG2354 Tropiocolotes cf. tripilotanus TG2354 0.97/54/100 Stenodactylus cf. sthenodactylus TG0185 Stenodactylus cf. sthenodactylus TG0185 Cyrtopodion scaber TG0109 Cyrtopodion scaber TG0109 0.96/71 Agamura persica TG0188 0.5/22/100 Agamura persica TG0188 Paroedura picta TG2139 Paroedura picta TG2139 Phelsuma laticauda TG1616 Phelsuma laticauda TG1616 C. boulengerii LSUHC 9278 C. boulengerii LSUHC 9278

0.85/61 C. psychedelica LSUHC 9244 Ca Mau C. psychedelica LSUHC 9244 C. limi LSUHC 3902 0.37/9/- C. limi LSUHC 3902 1.0/86 0.9/91/100 C. paripari LSUHC 9186 C. paripari LSUHC 9186 C. nigridia LSUHC 9170 C. nigridia LSUHC 9170 C. sundainsula MZBLace 10156 C. pemanggilensis LSUHC 8014 C. kendallii LSUHC 9178 C. rajabasa ENS 7766 C. rajabasa ENS 7766 0.89/0/94 Southern Sunda 0.92/0/- C. sundainsula MZBLace 10156 C. pemanggilensis LSUHC 8014 0.36/0/- C. kendallii LSUHC 9178 C. baueri LSUHC 7302 C. bidongensis LSUHC 11445 0.96/84 C. peninsularis HC 0228 1.0/90/- C. baueri LSUHC 7302 C. bidongensis LSUHC 11445 0.93/100/- C. mumpuniae MZBLace 10166 C. mumpuniae MZBLace 10166 1.0/100/- C. peninsularis HC 0228 1.0/100/- 1.0/78 C. argus LSUHC 10835 C. argus LSUHC 10835 C. flavigaster LSUHC 8835 C. flavigaster LSUHC 8835 C. karsticola LSUHC 9055 C. karsticola LSUHC 9055 0.88/3/100 C. perhentianensis LSUHC 8700 0.67/59/- C. perhentianensis LSUHC 8700 C. pseudomcguirei LSUHC 9145 0.77/11/100 C. grismeri LSUHC 9733

C. shahruli LSUHC 9163 0.76/98/- C. narathiwatensis USMHC 1350 C. mcguirei LSUHC 8853 C. flavolineata LSUHC 8079 C. grismeri LSUHC 9733 0.41/18/- C. stongensis LSUHC 11089 1.0/82 Malaysian C. mahsurae LSUHC11828 0.67/41/- C. pseudomcguirei LSUHC 9145 C. harimau LSUHC 9667 Peninsula 0.44/11/- C. hangus HC 225

0.12/87 C. affinis LSUHC 6759 0.94/19/- C. selamatkanmerapoh LSUHC 11015 C. flavolineata LSUHC 8079 C. temiah LSUHC 9817 0.58/62/- C. temiah LSUHC 9817 C. shahruli LSUHC 9163 0.7/93 35.3% C. narathiwatensis USMHC 1350 0.64/70/70 C. bayuensis LSUHC 9072 0.63/85 C. hangus HC 225 C. mcguirei LSUHC 8853 C. selamatkanmerapoh LSUHC 11015 0.53/87/- C. affinis LSUHC 6759 0.25/52 C. bayuensis LSUHC 9072 0.67/97/- C. mahsurae LSUHC 11828 C. stongensis LSUHC 11089 0.67/98/100 C. harimau LSUHC 9667 C. sp. nov. 1 LSUHC 9569 (MS 286) C. sp. nov. 1 LSUHC 9569 (MS 286) C. monachorum LSUHC 10807 C. monachorum LSUHC 10807 C. biocellata LSUHC 8789 Pattani C. biocellata LSUHC 8789 C. niyomwanae LSUHC 9568 C. niyomwanae LSUHC 9568 C. kumpoli LSUHC 8848 C. kumpoli LSUHC 8848 C. lineogularis ZMKU R 00728 C. caudanivea LSUHC 8577 C. caudanivea LSUHC 8577 37.3% C. chanthaburiensis LSUHC 9338 0.92/80/51 C. chanthaburiensis LSUHC 9338 C. lineogularis ZMKU R 00728 C. neangthyi LSUHC 8515 0.97/100/- C. neangthyi LSUHC 8515 0.88/98/- 0.98/78 C. aurantiacopes LSUHC 8612 C. tucdupensis LSUHC 8631 C. nuicamensis LSUHC 8649 0.5/27/- C. aurantiacopes LSUHC 8612 C. tucdupensis LSUHC 8631 0.84/99/- C. nuicamensis LSUHC 8649 C. phangngaensis BYU 62537 C. phangngaensis BYU 62537 Thai Peninsula 1.0/100/- C. chanardi LSUHC 9567 C. chanardi LSUHC 9567 C. sp. nov. 2 LSUHC 9570 (MS 283) Indochina C. sp. nov. 2 LSUHC 9570 (MS 283) 0.98/100/- 1.0/60 C. omari LSUHC 9978 C. omari LSUHC 9978 1.0/54 C. roticanai LSUHC11815 1.0/87/42 C. roticanai LSUHC 11815 C. punctatonuchalis LSUHC 12453 0.74/42/100 C. siamensis LSUHC 9474

C. huaseesom LSUHC 9458 0.25/1/- C. punctatonuchalis LSUHC 12453 C. siamensis LSUHC 9474 0.73/32/51 C. huaseesom LSUHC 9458 BS ≥90 and PP≥0.95 0.89/94 C. thachanaensis ZMKU R 00729 PP≥0.95, BS ≥70, and SVDBS ≥70 C. thachanaensis ZMKU R 00729 0.8 C. vandeventeri LSUHC 12449 0.8 C. vandeventeri LSUHC 12449

Figure
S3:
Phylogenomic
relationships
of
Cnemaspis
based
on
514
loci
(267,912
bp).
Major
clades
are
colored
based
on
the
distributions
used
in
Figure
1.
A.
Concatenated
ExaBayes
analysis
with
ExaBayes
PP
and
IQ- TREE
BS
support
values
respectively.
B.
Astral-II
species
tree
with
PP,
BS
(from
RAxML
bootstraps)
and
SVDBS
support
values
respectively.
The
numbers
in
bold
on
the
species
tree
represent
the
percentage
of
genetree
quartets
that
are
congruent
with
a
given
branch
(the
quartet
approach).
274 Correlophus ciliatus TG2314 Lialis burtonis TG2114 Coleonyx variegatus TG0179 100 Teratoscincus keyserlingii TG2382 Teratoscincus przewalskii TG233 100 100 100 Gonatodes ocellatus TG1855 100 Gonatodes vittatus TG1845 100 Sphaerodactylus macrolepis TG2147 100 100 Sphaerodactylus roosevelti TG2384 Sphaerodactylus townsendi TG2150 100 Phyllodactylus wirshingii TG2385 Thecadactylus rapicauda TG1864 100 Paroedura picta TG2139 100 Hemidactylus mabouia TG2142 Hemidactylus turcicus TG2125 100 100 100 Agamura persica TG0188 100 Cyrtopodion scaber TG0109 100 Stenodactylus cf. sthenodactylus TG0185 32 Tropiocolotes cf. tripilotanus TG2354 Gehyra mutilata LSUHC12328 Phelsuma laticauda TG1616 53 100 Cnemaspis boulengerii LSUHC 9278 Cnemaspis psychedelica LSUHC 9244 100 Cnemaspis limi LSUHC 3902 93 100 Cnemaspis nigridia LSUHC 9170 100 Cnemaspis paripari LSUHC 9186 Cnemaspis pemanggilensis LSUHC 8014 94 Cnemaspis sundainsula MZBLace 10156 100 Cnemaspis rajabasa ENS 7766 100 60 100 Cnemaspis kendallii LSUHC 9178 100 Cnemaspis baueri LSUHC 7302 98 Cnemaspis peninsularis HC 0228a 63 Cnemaspis bidongensis LSUHC 11445 Cnemaspis mumpuniae MZBLace 10166 GL115 98 Cnemaspis argus LSUHC 10835 100 99 Cnemaspis flavigaster LSUHC 8835 100 Cnemaspis karsticola LSUHC 9055 100 Cnemaspis perhentianensis LSUHC 8700 Cnemaspis grismeri LSUHC 9733 100 Cnemaspis stongensis LSUHC 11089 96 Cnemaspis flavolineata LSUHC 8079 76 Cnemaspis pseudomcguirei LSUHC 9145 62 45 Cnemaspis temiah LSUHC 9817 96 Cnemaspis bayuensis LSUHC 9072 85 Cnemaspis selamatkanmerapoh LSUHC 11015 70 60 Cnemaspis shahruli LSUHC 9163 97 Cnemaspis mcguirei LSUHC 8853 100 Cnemaspis harimau LSUHC 9667 Cnemaspis mahsurae LSUHC11828 100 Cnemaspis monachorum LSUHC 10807 100 Cnemaspis sp. nov. 1 LSUHC 9569 (MS 286) 100 Cnemaspis biocellata LSUHC 8789 100 Cnemaspis kumpoli LSUHC 8848 Cnemaspis niyomwanae LSUHC 9568 Cnemaspis caudanivea LSUHC 8577 51 99 Cnemaspis lineogularis ZMKU R 00728 98 100 Cnemaspis chanthaburiensis LSUHC 9338 90 Cnemaspis neangthyi LSUHC 8515 86 Cnemaspis nuicamensis LSUHC 8649 48 40 Cnemaspis aurantiacopes LSUHC 8612 Cnemaspis tucdupensis LSUHC 8631 Cnemaspis siamensis LSUHC 9474 Cnemaspis punctatonuchalis LSUHC 12453 42 51 100 Cnemaspis huaseesom LSUHC 9458 100 Cnemaspis thachanaensis ZMKU R 00729 34 Cnemaspis vandeventeri LSUHC 12449 98 Cnemaspis chanardi LSUHC 9567 100 Cnemaspis phangngaensis LSUHC 12438 100 Cnemaspis sp. nov. 2 LSUHC 9570 (MS 283) 99 Cnemaspis omari LSUHC 9978 Cnemaspis roticanai LSUHC11815

Figure S4: SVDQuartets consensus species tree with bootstrap consensus support values (SVDBS).

275 Correlophus ciliatus 45.7 Lialis burtonis Coleonyx variegatus Teratoscincus przewalskii 6.2 Teratoscincus keyserlingii † Gonatodes ocellatusz 77.3 21.0 105.9 Gonatodes vittatus Sphaerodactylus macrolepis 55.2 † 20.4 Sphaerodactylus townsendi 15.3 Sphaerodactylus roosevelti 94.9 Phyllodactylus wirshingii 64.3 Thecadactylus rapicauda Gehyra mutilata 73 Hemidactylus turcicus 19.8 Hemidactylus mabouia 84.5 Cyrtopodion scaber 62.8 26.1 Agamura persica 49.8 Tropiocolotes cf. tripilotanus 75.8 29.7 Stenodactylus cf. sthenodactylus Paroedura picta Phelsuma laticauda C. psychedelica 12.1 73.7 C. boulengerii C. limi C. paripari 21.8 69.7 5.9 C. nigridia 23.3 C. sundainsula C. kendallii 16.3 C. rajabasa 7.6 13.2 C. pemanggilensis 61.1 10 C. baueri C. peninsularis 6.9 4.0 C. bidongensis C. mumpuniae 3.1 C. argus C. flavigaster 16.4 C. perhentianensis 12.4 4.8 C. karsticola 41.9 C. pseudomcguirei 24.8 11.6 C. shahruli 6.7 C. grismeri C. mcguirei 14.8 C. affinis 10.4 5.2 C. mahsurae 12.4 3.2 C. harimau 9.3 C. temiah C. flavolineata C. narathiwatensis 11.3 C. selamatkanmerapoh 39.2 9.0 6.9 C. hangus 5.9 C. stongensis 3.4 C. bayuensis C. sp. nov. 1 22.1 10.6 C. monachorum C. biocellata

11.2 C. niyomwanae 5.9 C. kumpoli C. lineogularis C. caudanivea 20.5 37.3 C. chanthaburiensis 15.0 C. neangthyi 11.6 C. aurantiacopes 9.6 C. tucdupensis C. nuicamensis 33.4 5.4 C. punctatonuchalis C. huaseesom 18.7 C. siamensis 14.0 C. vandeventeri 11.8 C. thachanaensis 26.6 7.2 6.7 C. phangngaensis C. chanardi 8.7 C. sp. nov. 2

5.7 C. roticanai C. omari 5.0 Lower/Early Upper/Late Paleocene Eocene Oligocene Miocene Pli. P 110 100 80 60 40 20 0 Ma

Figure S5: Numbers near the nodes are mean divergence time estimates from fossil and root height calibrations implemented in MCMCtree. The cross symbols indicate outgroup nodes with calibration points. Plio=Pliocene, P= Pleistocene.

276 A. All Cnemaspis species B. No Ca Mau clade 10 10

8 8

6 6

4 4 speciation rate

2 2

0 0

70 60 50 40 30 20 10 0 Ma 70 60 50 40 30 20 10 0 Ma

C. All Cnemaspis species D. No Ca Mau clade 1.4 1.4

1.2 1.2

1.0 1.0

0.8 0.8

0.6 0.6

extinction rate 0.4 0.4

0.2 0.2

0.0 0.0

70 60 50 40 30 20 10 0 Ma 70 60 50 40 30 20 10 0 Ma time before present time before present

Figure S6: Estimated net speciation and extinction rates for Cnemaspis from BAMM outputs based on the time- calibrated tree. A. The rate shift for Cnemaspis estimated to be between 60–40 Ma with a higher magnitude of change from 2 to approximately 5.9 (=5.905598). B. The net speciation rate of Cnemaspis with the exclusion of the Ca Mau clade (the net speciation rate is constant =6.373306). C. Extinction rate for all of Cnemaspis with a small increase in extinction between 60–40 Ma (µ=0.41). D. Extinction rates with exclusion of the Ca Mau clade appear to be constant through time (µ=0.42).

277 Correlophus ciliatus Lialis burtonis Coleonyx variegatus Teratoscincus przewalskii Teratoscincus keyserlingii Gonatodes ocellatusz Gonatodes vittatus Sphaerodactylus macrolepis Sphaerodactylus townsendi Sphaerodactylus roosevelti Phyllodactylus wirshingii Thecadactylus rapicauda Gehyra mutilata Hemidactylus turcicus Hemidactylus mabouia Cyrtopodion scaber Agamura persica Tropiocolotes cf. tripilotanus Stenodactylus cf. sthenodactylus Paroedura picta Phelsuma laticauda C. psychedelica C. boulengerii C. limi C. paripari C. nigridia C. sundainsula C. kendallii C. rajabasa C. pemanggilensis C. baueri C. peninsularis C. bidongensis C. mumpuniae C. argus C. flavigaster C. perhentianensis C. karsticola C. pseudomcguirei C. shahruli C. grismeri C. mcguirei C. affinis C. mahsurae C. harimau C. temiah C. flavolineata C. narathiwatensis C. selamatkanmerapoh C. hangus C. stongensis C. bayuensis C. sp. nov. 1 C. monachorum C. biocellata C. niyomwanae C. kumpoli C. lineogularis C. caudanivea C. chanthaburiensis C. neangthyi C. aurantiacopes C. tucdupensis C. nuicamensis C. punctatonuchalis C. huaseesom C. siamensis C. vandeventeri C. thachanaensis C. phangngaensis C. chanardi C. sp. nov. 2 C. roticanai C. omari Lower/Early Upper/Late Paleocene Eocene Oligocene Miocene Pli. P 100 80 60 40 20 0 Ma Figure S7: The BAMM tree with a single rate shift (blue circle, probability=0.70) associated with a rapid radiation of
Cnemaspis
with
the
exclusion
of
the
Ca
Mau
clade. 278 iueS:A ieg hog iepo ae ntetm-airtdte eeae rmMMte.B Null B. observed MCMCtree. the with from MCCRT generated line[7]. the ( black tree from tree solid trees time-calibrated empirical vertical rate the the constant on for pure-birth statistic based simulated gamma 20,000 plot the time from through distribution Lineage gamma A. S8: Figure

number of spp. A. 0 10 20 30 40 50 05 03 01 0 10 20 30 40 50 60 Time 1871,pvle=001 rtclvle=-.5)rpeetda the as represented -1.758) = value critical 0.041, = p-value -1.847611, = B. Frequency 0 1000 2000 3000 4000 4- 2 0 -2 -4 null gamadistribution 4 279 A B C Frequency Frequency Frequency 0 10 20 30 40 50 0 10 20 30 40 0 10 20 30 40

10 20 30 40 50 60 70 200 300 400 500 600 700 800 0 50 100 150 200 Number of Taxa Alignment length (bp) Number of Informative Sites

D E F R2= 0.0854 p <0.0000 Frequency Frequency Informative Sites Informative 0 50 100 150 200 0 10 20 30 40 50 0 50 100 150

0 10 20 30 40 0 5 10 15 20 25 30 35 200 300 400 500 600 700 800 Percentage of Informative Sites Missing data (%) gaps &N’s Alignment Length

Figure S9: Summary statistics from the raw Illumina reads. A. Number of taxa in the alignment. B. Alignment length C. The number of informative sites. D. Percentage of informative sites. E. The percentage of missing data, gaps and N’s. F. Alignment length with respect to informative sites; there is a significant positive correlation between the alignment length and the number of informative sites.

280 References

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[3] Hall, R. The palaeogeography of sundaland and wallacea since the late jurassic. Journal of Limnology 72,e1 (2013).

[4] Grismer, L. L. et al. Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa 3880, 1–147 (2014).

[5] Minh, Q., Nguyen, M. & von Haeseler, A. A. Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30, 1188–1195 (2013).

[6] Nguyen, L.-T., Schmidt, H. A., von Haeseler, A. & Minh, B. Q. IQ-TREE: A fast and e↵ective stochas- tic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32, 268–274 (2015).

[7] Pybus, O. G. & Harvey, P. H. Testing macro–evolutionary models using incomplete molecular phylogenies. Proceedings of the Royal Society of London. Series B: Biological Sciences 267, 2267–2272 (2000).

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